Ending with Elton: Preludes to Invasion Biology

©2006 PLEASE FORWARD ALL DISTRIBUTION REQUESTS TO THE AUTHOR ENDING WITH ELTON PRELUDES TO INVASION BIOLOGY by Matthew K. Chew A Dissertation Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy Arizona State University December 2006 ©2006 PLEASE FORWARD ALL DISTRIBUTION REQUESTS TO THE AUTHOR ENDING WITH ELTON PRELUDES TO INVASION BIOLOGY by Matthew K. Chew has been approved November 2006 APPROVED: Manfred D. Laubichler, Chair Jane Maienschein James P. Collins George Thomas Gregg Mittman (by A.L Hamilton) Supervisory Committee ACCEPTED: Miles Orchinik Director of the School Maria T. Allison Dean, Division of Graduate Studies ©2006 PLEASE FORWARD ALL DISTRIBUTION REQUESTS TO THE AUTHOR ABSTRACT Invasion Biology, the study of biota redistributed via human agency, has traditionally traced its founding to Charles Elton’s 1958 book The Ecology of invasions by Animals and Plants. But there were many substantial, scientific preEltonian accounts and analyses of redistributed biota dating back at least to the mid-1700s; and non-Eltonian treatments appeared into the late 1950s. Elton began writing on the topic by 1925. From 1931 to 1948 he developed his ideas on conservation in association with Aldo Leopold. Their “competitive collaboration” is explored and documented, showing that each supported and contextualized the other. Elton was, in part, inspired to write The Ecology of invasions by Animals and Plants in dissatisfied response to an earlier effort by American Marston Bates. The two authors, and these works, are compared and contrasted. The Ecology of Invasions by Animals is analyzed in detail, showing how Elton framed his arguments in terms of warfare and cold-war era nuclear alarmism, making a plea for action without providing a strong theoretical basis for preventing the redistribution of biota. The republication of Elton’s book in 2000 incited a new round of interpretation, but even while its proponents hailed the book as invasion biology’s ‘bible,’ they prepared to replace it with more modern texts. The narrative shows how a ideas about the origins of life on earth, the role of humans in nature, a sense of place and biogeographical belonging, and concerns about the unintended consequences of human agency motivated scientists to attempt to impute order to, and impose it upon, the historically contingent distribution of biota. ©2006 PLEASE FORWARD ALL DISTRIBUTION REQUESTS TO THE AUTHOR To a very patient Juliet. “On entering this task I did not realise how vast it was, and how fragmentary was the sum of existing knowledge, but having commenced it, I had no thought of turning back, or of abandoning the project. Even if the record be imperfect, it will be of some use to future workers to have pieced together the available material.” — George M. Thomson, The Naturalisation of Animals & Plants in New Zealand iv TABLE OF CONTENTS Page LIST OF FIGURES............................................................................................... vi INTRODUCTION...................................................................................................1 CHAPTER 1. SCIENCE IN A FUGUE STATE.....................................................11 2. THE MUTUAL ADMIRATION OF CHARLES ELTON AND ALDO LEOPOLD........................................................91 3. WAR AND PEACE: VIEWS OF BIOTIC REDISTRIBUTION IN THE 1950s ...................................................................175 4. THE ECOLOGY OF INVASIONS BY ANIMALS AND PLANTS, A SUMMARY REVIEW .....................................................208 5. ELTON AND INVASIONS, RECONSTITUTED AND RETIRED..................................................................269 GENERAL CONCLUSIONS..............................................................................283 REFERENCES..................................................................................................289 2 LIST OF FIGURES Figure Page 4.1 Advertisement for The Ecology of Invasions, 1958 ....................................211 4.2 Bur oak acorns ...........................................................................................231 4.3 Grandibus exigui sunt pisces piscibus esca ...............................................257 4.4.a. The German invasion of France.............................................................255 b. Breeding range of the European starling in eastern North America .......256 c. The German catastrophe in White Russia..............................................257 iv Introduction: Science in a Fugue State In 1999, marine ecologist James T. Carlton enlisted as the founding editor of the journal Biological Invasions. His inaugural editorial explained the journal’s topical focus; a study its participants called invasion biology: Tens of thousands (or perhaps hundreds of thousands) of species of bacteria, protists, fungi, animals, and plants have been rapidly transported–and successfully established– due to human aided movement across and between continents and oceans. This ‘synanthropic’ process is not simply a matter of ‘speeding up what would happen anyway’: an enormous number of these species have achieved new distributions that they would never have gained (at least in their present form) over geological time. As a result of instantaneous human-assisted transport, species are abruptly faced with interacting with new species and new environments with which they have had no evolutionary history. At least partially as a result of this, a very large number of these invasions have caused extraordinary environmental, social, and economic changes. The Earth is now virtually itching with new invasions at, one might argue, an unprecedented rate if taken in a global context. (…) And yet, surprisingly, despite the scales of natural and human-mediated invasions, no international platform has ever existed for scholarly communication devoted to these phenomena that have changed, and now are changing, the face of the Earth so dramatically.1 Over the next few years, invasion biology’s frankly alarmist claims became increasingly common in news and documentary coverage, particularly in Anglophone countries. Governments responded. In the United States, invasion biology became a charter function of the new Department of Homeland Security. Where did this new, publicity-prone kind of biology come from? Other arguable “foundings” of invasion biology occurred in the 1980s and 1990s. Notable among them was a proposal spearheaded by South African scientists, 1 James T. Carlton, "Editorial: A Journal of Biological Invasions." Biological Invasions 1 (1999) 1. 2 culminating in a 1982 declaration by the Scientific Committee on Problems of the Environment (SCOPE) that “the invasive spread of plants, animals, and microorganisms introduced by man into areas remote from their centres of origin” constituted an “environmental problem.”2 That action prompted a spate of regional symposia and published proceedings. Finally, in 1998 a review article in BioScience, the journal of the American Institute of Biological Sciences, reified and expanded a 1992 assertion by Edward O. Wilson that “invasive exotic species are the second greatest threat to biodiversity.”3 But invasion biology’s devotees frequently memorialized an origin some forty years prior to the first issue of Carlton’s journal. “As much as one can ever pinpoint the beginning of a field to a single event, the [1958] publication of Charles S. Elton’s The Ecology of Invasions by Animals and Plants signaled the beginning of the field of invasion biology,” wrote a reviewer celebrating the book’s 2000 reissue.4 In his new foreword to an otherwise unaltered facsimile edition, Daniel Simberloff wrote that Elton’s book “founded a whole field of research,” and “was to become a bible for practitioners of a burgeoning new science.”5 2 I.A.W. Macdonald, F.J. Kruger and A.A. Ferrar, The Ecology and Management of Biological Invasions in Southern Africa (Capetown: Oxford University Press, 1986), v. 3 David Wilcove, et al, “Quantifying Threats to Imperiled Species in the United States” BioScience 48 (1998) 607-615; Edward O. Wilson, The Diversity of Life (Cambridge, MA: Belknap, 1992) 253-4. 4 Ingrid M. Parker, “Invasion Ecology: Echoes of Elton in the Twenty-First Century” Conservation Biology 15 (2001) 806-807. 5 Daniel Simberloff, “Foreword” IN Charles S. Elton, The Ecology of Invasions by Animals and Plants. (Chicago: University of Chicago Press, 2000) vii. iv 3 Harold A. Mooney, Executive Committee Chairman of the self-styled Global Invasive Species Programme (GISP) asked science writer Yvonne Baskin to write a book about invasion biology as “part of the GISP effort.”6 She was “granted access to worldwide science and policy discussions of [GISP] through one of its sponsors, the Scientific Committee on Problems of the Environment [SCOPE],” suggesting that her volume would stand as the public apologia and authorized history of the field. As she represented the collective conception of GISP and SCOPE, “the first person to … lay the foundation for a science of invasion biology was pioneering British ecologist Charles Elton [who] first pulled together his thoughts on bioinvasions [sic] … in 1957.”7 Notwithstanding some debatable terminology, the leaders of a movement calling itself invasion biology considered Charles Elton’s 1958 monograph to be the seminal, indispensable discussion of the phenomena that preoccupied them. Furthermore, they claimed that Elton himself had not substantially addressed the matter previously. But Elton’s book was obviously a review, not a revelation. Its fourteen-page bibliography included scientific accounts published as little as a year and as much as a century prior. All of its line illustrations and most of its plates were reproduced from earlier works by other authors. Invasion biology had a history to explore; 1957 was no more satisfactory than 4004 B.C. 6 Yvonne Baskin, A Plague of Rats and Rubbervines: The Growing Threat of Species Invasions. (Washington DC: Island Press, 2002) 8. 7 Ibid.,8-9, 128 and first edition dust jacket. iv 4 This dissertation rediscovers and relates the history of scientific investigation and discussion of biota redistributed via human agency (neobiota, following). 8 Responding to the claims of Baskin and Simberloff, my thesis has three parts: (1) Scientists began observing and theorizing about neobiota no later than the mid-eighteenth century. (2) Charles Elton “pulled his thoughts together” regarding neobiota episodically, and published them several times, beginning in the 1920s. (3) Invasion biologists and their allies looked no further than Elton and The Ecology of Invasions by Animals and Plants for a “bible” and a founder because the erewhile celebrated ecologist’s alarmist book provided authoritative support for their own alarmist responses to neobiota. The Purposes of this Study It is important for invasion biologists to understand that neither their interests nor their concerns arose de novo in the 1950s, much less the 1980s or 1990s. That belief unnecessarily isolates them from a diverse array of prior work, in some cases by scientists revered before and beyond Charles Elton. To paraphrase Santayana, invasion biologists who remain ignorant of centuries of prior work are condemned (if lucky) to repeat it, or (if unlucky) to simply remain ignorant. Historians of science will find here another expression of some familiar 8 The convenient and neutral shorthand neobiota is common in recent European literature, particularly in German language publications. iv 5 themes, but also a new emphasis on the development of attempts to normatively relate, and even attach, biota to places. Charles Elton’s celebrated concern over neobiota was not the mature judgment of a veteran ecologist. It was the reflexive response of a young man, perhaps even a child, doting on and deferring to a brilliant, authoritarian elder brother through a wartime adolescence. As the decades passed Elton’s exposition and justifications grew more sophisticated, but not his attitude. This narrative reveals that process and re-humanizes the hero, not to discard him, but to remind his aspiring emulators and followers that not everything a brilliant scientist has to say is necessarily brilliant or scientific. Few of the expert opinions about neobiota that preceded Elton’s were alarmist, and those were not all alarmist for Elton’s reasons. Invasion biology’s selection of The Ecology of Invasions as its “bible,” in light of both a completely new analysis of the book, and a review of its previous reviewers, shows the differences and similarities between Elton’s meaning and the meaning invasion biology gave him. This instance of investing contemporary interests with the perceived authority of a dead celebrity reminds us all just how easily influenced even scientists are by both nostalgia and “star power.” iv 6 The Containment Strategy To support my thesis and elaborate the points just made, this study offers a cast of characters drawn mostly from Enlightenment and post-Enlightenment Europe, including expatriates and their descendants, bearing the intellectual stamps of European science and colonialism. In order to make this a history of science, the authors discussed here were all “doing science” according to the standards of their times, acknowledged as competent (or at least, not dismissed as incompetent) by their peers. By that criterion, many non-scientists and some “near-scientists” have also commented on neobiota, often astutely, but they must await future work. In the course of developing narratives that rather unexpectedly grew to span a quarter millennium of global activity, I encountered many potentially fruitful avenues of investigation and analysis. As the need arose I made brief sorties along these “roads not taken,” usually to contextualize a historical figure, or to provide preliminary framing of tangential and subsidiary questions that will require a career to answer fully. As one example, I could not describe The Ecology of Invasions or discuss invasion biology without addressing the trope of “invasion”. Elton’s book is shot through with military imagery. Larger questions loomed regarding the role of figurative language in biological sciences, but I did not review that literature or attempt to generalize in that regard.9 9 Manfred Laubichler and I commented briefly on biological metaphors in “Natural Enemies: Metaphor or Misconception?” Science 301 (2003) 52-53; Brendon M.H. Larson and colleagues discussed the militarization of conservation in “The iv 7 Among other potential themes arising, the status of both invasion biology and its preludes as scientific disciplines is debatable, but undertaking that analysis would overcomplicate this general history. European colonialism and its aftermath could also have become a major preoccupation, but like the other potential frameworks mentioned here, it made only cameo appearances. The relationship between science and religious belief is a recurring feature of this narrative, but making it a major analytical theme is a task for another volume. The disparaging epithets sometimes applied to neobiota by invasion biologists and their allies have been compared to racism by authors from various disciplines.10 I did not elide this problem when it arose in the material under examination, but neither did I pursue a comprehensive analysis. Overall, the strategy was to make my necessary major points while acknowledging, but not indulging, the many tempting tangents. Sources The material analyzed and presented here was gathered from both published and unpublished sources. Except as noted, all quotations retain their original emphasis. Most were written in English, and all but a few of the others War of the Roses: Demilitarizing Invasion Biology” Frontiers in Ecology and the Environment 3 (2005) 495-500, and “Metaphors and Biorisks: The War on Infectious Diseases and Invasive Species.” Science Communication 26 (2005):243-268.. 10 See, e.g., Ned Hettinger, 2001. “Exotic Species, Naturalisation, and Biological Nativism.” Environmental Values 10:193-224; Banu Subramaniam, “The Aliens Have Landed! Reflections on the Rhetoric of Biological Invasions.” Meridians: feminism, race, transnationalism 2 (2001) 26-40; David Theodoropoulos, Invasion Biology: Critique of a Pseudoscience (Blythe, CA: Avvar, 2003). iv 8 were quoted from published English translations. In the rare cases where no published translation of an essential document was available, I identified and translated the relevant passages. Chapters Chapter One, “Biota and Belonging Before 1950,” demonstrates that there were many substantial, scientific pre-Eltonian accounts and analyses of redistributed biota dating back at least to the mid-1700s; and non-Eltonian treatments appeared into the late 1950s. Working within several different traditions and milieux, most of these authors encountered and considered redistributed biota as it impinged upon their main interests. Some of their names are familiar because of their contributions within those contexts. Others have been largely forgotten, except perhaps by taxonomists who memorialize their authority, if not necessarily their contributions. In another effort at topical containment, I did not elucidate the allied tradition of agricultural pest suppression, because its objectives do not vary significantly with the geographical origin of the target species.11 Charles Elton began “pulling his thoughts together” on the topic at hand long before 1957. Chapter Two, “The Mutual Admiration of Charles Elton and 11 See, e.g., George Ordish, The Constant Pest (New York: Scribners, 1976) and Mark L. Winston, Nature Wars: People vs. Pests (Cambridge: Harvard University Press, 1997). Philip J. Pauly wrote two interesting case studies touching on the matter, "The Beauty and Menace of the Japanese Cherry Trees: Conflicting Visions of American Ecological Independence." Isis 87 (1996) 51-73; and "Fighting the Hessian Fly: American and British Responses to Insect Invasion 1776-1789." Environmental History 7 (2002) 378-99 iv 9 Aldo Leopold,” reviews Elton’s relevant, earlier published works, and examines his available unpublished writings and correspondence. It follows the development of his ideas and attitudes about “invasions” and provides enough biographical context to identify his major influences. It demonstrates that Elton’s general feelings about invasions dated back at least to his student years at Oxford, and that over time he became more vehement, rather than more circumspect, in expressing them. Elton emerges as an interesting if still enigmatic figure, and his “competitive collaboration” and friendship with Aldo Leopold is shown to have been a crucible of both “invasion” dogma and conservation philosophy and policy. Chapter Three, “War and Peace: Views of Neobiota in the 1950s,” shows that non-Eltonian views persisted until Elton’s book appeared. It describes the relationship between “Balance and Barrier,” Elton’s proto-Invasions BBC radio lecture series on conservation, and another synthesis, “Man as an Agent in the Spread of Organisms” by American animal ecologist Marston Bates. It compares and contrasts the two authors and the two works, showing that the Bates paper exemplified a more objective approach to the topic than Elton’s. It describes the interactions between these scientists, and suggests that even though Elton did not explicitly engage with Bates’s analysis, his dissatisfaction with “Man as an Agent” was a proximate motivator for writing The Ecology of Invasions. Chapter Four, “Dissecting The Ecology of Invasions by Animals and Plants,” describes the book’s development from conception through early reviews iv 10 and citations. Building on the context established by the previous chapters, it shows to the extent possible what and how Elton was thinking, what he meant, whom he meant it for, how he said it, and why. It explores his use of visuals, metaphors and literary allusions. It shows that his scientific purpose in writing was secondary to his socio-political purpose: to inoculate the public with his own disquiet — his “fear and loathing” — of “invasions by animals and plants;” and further, to persuade them of the merits of his own conservation philosophy. Chapter Five, “Elton and Invasions, Reconstituted and Retired” is a relatively brief foray beyond 1958, examining the 2000 reissue of The Ecology of Invasions and the way invasion biologists then reframed it to suit their own purposes. It shows how the edition’s new introduction by Daniel Simberloff (the first insertion of non-Eltonian material) not only hailed the book as invasion biology’s bible, but served as that bible’s exegesis, a guide for reading Elton’s idiosyncratic jeremiad as revelation — in the “proper” light of later developments. Motivations As an ecologist, conservation professional, and (now) nascent historian of biology, I intended the products of this project to appeal, or at least, speak, primarily to members of those three disciplines. But it should engage anyone with a contemplative interest in nature, including human nature. I wrote it as a doctoral dissertation explicitly to bring external discipline and merit review to bear on what might otherwise evolve into an outré polemic of my own. iv 11 My first fifteen years as a student and practitioner in those fields coincided with the emergence of invasion biology. During that period I earned two degrees, conducted endangered species and breeding bird surveys and field research in riparian ecology, assisted in National Park visitor perceptions research, wrote and co-authored interpretive materials, environmental documentation, peerreviewed research papers, book chapters and policy documents. I donned protective gear and a backpack full of 2,4 D and resisted, to little avail, the advent of Euphorbia esula along a northern Colorado floodplain. I worked as an ecological consultant, and then as the “coordinator” of two state conservation programs, usually in close cooperation with major conservation charities. I served on the board of a statewide conservation organization and on the “education and outreach” committee of an international, professional one. In short, as both a producer and user of information during that period I had a pretty good view of the ongoing action. What I saw of the products and practice of invasion biology looked more like propagating and defending dogma than producing and disseminating knowledge. I wanted to understand why. What follows is part of what I learned. iv Chapter 1 Biota and Belonging Before 1950 There are many ways to divide invasion biology’s preludes, none of which are self-evidently satisfactory. Like a genealogy, they only amount to a “whole” with respect to one confluence, and can only be seen as such retrospectively, from there. For the purposes of this discussion, that point is Charles Elton’s 1958 book, The Ecology of Invasions by Animals and Plants. The authors and works discussed here are the major scientific precursors of Elton’s 1958 synthesis. He cited a few, and knew of a few more, but was apparently unaware of most of them. Sorting these works into meaningful subgroups for analysis was not a straightforward task. Their phylogeny was complicated by both crossfertilizations and (now) odd-seeming self-sortings. Even defaulting to dividing them into botanists and zoologists failed to make complete sense. But my first cut divides these earlier workers into two camps, based on a few general aspects inherent in the features of the biota they emphasize. Members of one camp studied taxa that were readily collectible, preservable, and included a large yet finite number of identifiable, regionally familiar taxa. Vascular plants, shelled mollusks and butterflies were all collected, pored over, exchanged, and written up by cadres of (often amateur) natural history enthusiasts. By the early 1800s comprehensive inventories of these taxa were being assembled in most European countries and in populated regions of their temperate colonies. This established a baseline for comparison, allowing 13 anomalies to be quickly discerned and perhaps recognized as new arrivals. One particularly strong tradition developed among botanists, and will be discussed at some length. Conchologists are covered more briefly, but appear likely to offer fertile ground for additional research. Neobiotic butterflies and (particularly) moths were usually covered in the literature of agricultural pest management, which I have chosen not to consider here (see introduction). All other taxa default to the rather amorphous super-category of unmet conditions: e.g., cryptogamous plants, motile animals, anything microscopic, inaccessible, nocturnal, fossorial or rare, pelagic, benthic, indistinguishable or otherwise hard to observe and identify. Specialists dealing with taxa exhibiting such characteristics were mostly professional scientists, explorers, or wellfinanced, advanced amateurs, often working in relative isolation and developing their own methods. This chapter is organized by traditions (where discernable), taxa under consideration, and geography; and within those divisions, by author and chronology. All fit into a very large story of the motivation and development of biogeography. But since my purpose is mainly to demonstrate the content and pre-Eltonian existence of a particular sort of literature, I will only revisit the elements of that large story that are most germane to studying neobiota. Botany and Cameralism: Peter Kalm The age of European exploration and colonization was fraught with transoceanic and transcontinental introductions. Colonists (“beginning” with Columbus’s second voyage in 1493) carried the familiar materia agricola of iv 14 subsistence farming with them, and merchant-explorers returned with pricey exotica such as fruits, seeds and spices. Competition between countries and companies to locate sources and monopolize such products was fierce. Finding ways to produce them closer to home was a priority for countries with few colonial claims. Without claiming it categorically as “the first scientific account” of introduced biota, one early publication stands out enough to make it worth detailing. On 30 November 1747, Peter (Pehr) Kalm departed from Gothenburg, Sweden on a voyage of horticultural espionage.12 Writing in the 1970s, Frank Egerton and Ralph Sargent each cast Peter Kalm in the role of America’s first, or best, early ecologist.13 Kalm’s foray into the wider world was long planned and his objective had always been to identify and gather up useful foreign plants for acclimatization and exploitation in Sweden. His mentor, Carl Linnaeus believed that America offered the best hope of harboring unknown plant species that could be cheaply and quickly collected, then commercially grown in Sweden, thus quickly transforming the kingdom’s then-diminishing fortunes and making it an economically independent powerhouse. 12 Martti Kerkkonen, Peter Kalm's North American Journey: Its Ideological Background and Results, Studia Historica. (Helsinki: Finnish Historical Society, 1959); Lisbet Koerner, “Linnaeus' Floral Transplants.” Representations 47: 144169. Christened Petter by his Finnish parents, Kalm was later referred to as Pehr in Sweden and as Peter and Pierre respectively by English and French speaking acquaintances during his travels (Kerkkonen, Kalm’s … Journey, 47). 13 Ralph Sargent, Introduction, IN Peter Kalm, Travels into North America, trans. J.R. Forster (Barre, MA: Imprint Society, 1972) xxiv; Frank N. Egerton, “Ecological Studies and Observations before 1900” IN Issues and Ideas in America, ed. B.J. Taylor and T.J. White (Norman: University of Oklahoma Press, 1976), 311-51. iv 15 Kalm’s departure was repeatedly delayed. In an April 1746 letter to Pehr Elvius, secretary of the Swedish Academy of Sciences, Kalm professed his frustrated eagerness to be off: “Good Sir, my youth is passing; this is the time when everything delights the eye and nothing so rejoices the mind as the contemplation and collecting of things in Nature; an unknown herb, an insect that I have not seen before, a new use for some natural product, can give me more pleasure than the discovery of a treasure gives a miser.”14 Peter Kalm did not set out to find European plants in America, but his journal, especially for the first year of his visit, confidently identified fifteen familiar Swedish species. Some he thought had obviously been planted on purpose and spread or persisted after cultivation: hemp, privet, and English ivy. He saw uncultivated parsnips “every day” during his wanderings in New York, and assumed them to be the garden variety; but they could have been accidentally introduced “wild” parsnips of the same species but lesser roots. Other introductions were most likely accidental, but he did not speculate about this phenomenon. Many remain familiar to North American gardeners, notably including dandelions, purslane, white clover and broad plantain. Several more plants that could easily have been indigenous but have strong resemblances to European relatives were too vaguely discussed to be useful here. He saw “red clover”, but there were red-flowered clovers on both sides of the Atlantic; and “common garden cresses” that could just as easily have been American peppergrass. 14 Kerkkonen, Kalm’s … Journey (cit. n. 12), 69. iv 16 Kalm was hardly an entomologist, but mostly to his dismay and discomfort he recognized a few European insects. Bedbugs were a common problem for colonists, but supposedly not for native Americans. He was told that houseflies had arrived some 150 years earlier, and did not doubt that those he saw were of European extraction. One of his longer notations concerns the “plague” of “mill beetles” (cockroaches). Dr. Cadwallader Colden, who claimed to have personally witnessed the ongoing seaborne arrival of the insects via ships from Caribbean islands, called cockroaches “properly natives of the West Indies.” Kalm found that claim plausible regarding urban roaches, but rhetorically asked “how can that be said of those mill-beetles, which are found in the midst of woods and deserts [wilderness]?”15 He more happily discussed honeybees, relating in perhaps his bestremembered anecdote that “The Indians… generally declare, that their fathers had never seen any bees either in the woods or any where else before the Europeans had been several years settled here [the Philadelphia area],” and that in acknowledgment the natives named honeybees “English flies.” He also noted that honeybee swarms were generally thought to fly only south, presumably seeking milder climes.16 Kalm’s notes suggested other establishments-in-progress. He saw “rats” without differentiating between types, assuming those found around settlements 15 Peter Kalm, Travels into North America (cit. n. 13), 208. Colden, a Scot by birth, was a major figure in colonial New York; he was also a sympathetic authority on the Native Americans of the region and likely facilitated Kalm’s contacts with them. 16 Ibid.,149. iv 17 to be European, but noting that John Bartram told him native rats could be found in regions yet unsettled.17 He assumed that the mice he saw were common house mice. He was told that red foxes were a recent addition to New England, but correctly doubted the veracity of that claim. Kalm may have described another insect arrival without realizing it. His entry for 1 October 1748 discussed mosquitoes (which he freely referred to as “gnats” but specifically identified as Culex pipiens), comparing the relative severity of bites he received in New Jersey and Sweden. According to “the Old Swedes” living along the river Morris, “the gnats had formerly been more numerous; that even at present they swarmed in vast quantities near the sea shore, near the salt water; and that those that troubled us this autumn in Philadelphia were of a more venomous kind than they commonly used to be,” judging by the greater distress resulting from their bites. Giving us a window into the proprieties of being a Swedish gentleman, Kalm professed that “when they stung me a night, my face was so disfigured by little red spots and blisters, that I was almost ashamed to shew myself.”18 The changes in both population densities and effects attributed to the mosquitoes around Philadelphia could easily have heralded the arrival of a new species. Kalm was considered a keen observer and a quick study. He declared himself an ideal empiricist: “I have made it a rule for myself that I don’t hold anything certain in natural history until I have seen it and carefully examined it 17 Ibid.,.225. 18 Ibid., 79-80. iv 19 myself.” 18 He was an inveterate note-taker who left a substantial documentary legacy. His detailed account of the North American mission was published in the form of an intermittent journal. It ran to multiple volumes, only two of which were published in Sweden, translated into German, and then finally from German to English by another notable botanical voyager, Johann Reinhold Forster. Unfortunately, publication was never completed, and much of the unpublished material was presumed destroyed in an 1827 library fire.20 Neobiota and the Beginnings of Biogeography Many monumental works are treated very briefly in this section, and my tiny excerpts and summary statements provide nothing like a general overview of any of them. But they do suggest each author’s awareness of and attitude toward the human role in biotic distribution. George Louis LeClerc, comte de Buffon Buffon’s Histoire Naturelle was a prime and seminal example of an Enlightenment era attempt to describe, if not necessarily explain, the earth’s diversity. Buffon recounted various second-hand tales of species introduction, e.g.: rats from Europe to various colonies; goats, sheep and mice from Europe to America; ferrets from Africa to Spain; and dogs to Africa. 21 But his particular contribution was more general: “That different regions of the globe are inhabited 19 Kerkkonen, Peter Kalm’s … Journey (cit. n. 12), 56. 20 Sargent, Introduction to Kalm, Travels (cit. n. 13). 21 Georges Louis Leclerc, comte de Buffon, The natural history of animals, vegetables, and minerals; with the theory of the earth in general. Transl. W. Kenrick and J. Murdoch. (London: T.Bell 1775-1776) Volume 2: 124, 128, 213, 336, 402. iv 19 by entirely distinct animals and plants is a fact which has been familiar to all naturalists since Buffon first pointed out the want of specific identity between the land quadrupeds of America and those of the Old World.”22 Karl Willdenow Pharmacist Karl Ludwig Willdenow was named Director of the Berlin Botanical Garden in 1801, but by then had already compiled a massive personal herbarium and published two major botanical works. His 1792 Grundriss der Kräuterkunde zu Vorlesungen was translated into English in 1805, retitled Principles of Botany. Willdenow was a very early phytogeographical theorist who (mostly on the basis of received specimens) developed elementary ideas, such as that similar latitudes often supported similar plants, but similar longitudes did not.23 Before receiving many New World specimens, he expected that (all else being equal) one would “meet in America, Asia and Africa, in plains of the same latitude, plants in common to the three parts of our globe. He hypothesized that “plants are disseminated from the highest mountains toward the flat countries” resulting in “five principal floras in Europe.”24 With regard to neobiota, Willdenow opined, “I think there is hardly one plant which originally grew wild in all latitudes. Plants, which are thus far 22 Charles Lyell, Principles of Geology Vol. 2. (London: John Murray, 1832) 66. However, Buffon did not discount the possibility that one set derived from the other, and “degenerated” into different species. He even proposed that presently separated continents were once united. 23 Karl Ludwig Willdenow, Principles of Botany, trans. anon. (Edinburgh: William Blackwood, 1805) 373. 24 Ibid., 402. iv 20 disseminated, were so by man.” Even though he immediately suggested that some commonly known examples of multi-latitude species were probably congeners mistaken for one another, he finished his discussion of “the history of plants” on a curious note: “I do not doubt, however, that of so vast a number of plants, there may be some which have so favourable a constitution, as to bear all climates; as in the animal kingdom, man, dogs, and swine, which agree with every possible climate.”25 Much of Willdenow’s phytogeography was shortly to be overturned, but he had provided testable hypotheses. Alexander von Humboldt In addition to running an apothecary shop, Karl Willdenow provided botanical lectures and field trips for interested Berliners. His best-remembered protégé was Alexander von Humboldt, who went on to conduct the first largescale studies of phytogeographical phenomena, describing the dependence of vegetation on abiotic factors such as climate and soil composition, and contributed significantly to describing the biogeographic analogy between increasing latitude and increasing elevation. Humboldt’s life is relatively welldocumented and can be treated briefly here. Like Buffon’s and Willdenow’s, his contribution to the study of neobiota was a largely theoretical one, relating plants to the places and conditions they grow under. But in 1805 he remarked on the commonness of their (agricultural) dispersal by humans: “it is not only winds, currents and birds that aid the migration of plants; man primarily takes care of 25 Ibid., 408. iv this.” 26 21 Humboldt later mentioned the establishment of American cacti in the Mediterranean region: “Cactus opuntia has spread during the last quarter of a century in a remarkable manner through Northern Africa, Syria, Greece, and the whole of southern Europe; penetrating the coasts of Africa far into the interior, where it associates with the native plants.”27 Augustin Pyramus (A.P.) de Candolle28 Among Humboldt’s eventual friend-correspondent-collaborators was Swiss-born botanist Augustin Pyramus de Candolle, whose career is relatively well-documented and will not be recounted in detail here. Among his many contributions, he coined the term “taxonomy” in his 1813 Théorie Élémentaire de la Botanique.29 de Candolle wrote this book while working in Montpellier, France, and in it mentioned the abundance of foreign plants established there.30 Later, writing nearly thirty years after Willdenow and benefiting from Humboldt’s (and other interim) discoveries, A.P. de Candolle nudged phytogeography toward a more empirical underpinning. In 1820 he published Essai Élémentaire de 26 Alexander von Humboldt, “Essai sur la Géographie des Plantes,” trans. Francesca Kern and Phillipe Janvier in Mark V. Lomolino, et al, Eds. Foundations of Biogeography (Chicago, University of Chicago Press 2004) 49-57. 27 Alexander von Humboldt, Views of Nature: or Contemplations on the sublime phenomena of creation; with scientific illustrations. trans. E. C. Otté and Henry G. Bohn (London: Bohn 1850). 28 I have seen “de Candolle” rendered “De Candolle,” and “Decandolle,” and sometimes two different ways in the one article. Modern libraries tend to catalog the name as “Candolle.” 29 Oxford English Dictionary Online, accessed 11 October 2006. 30 Viktor Mühlenbach, "Contributions to the Synanthropic (Adventive) Flora of the Railroads in St. Louis, Missouri, U.S.A." Annals of the Missouri Botanical Garden 66.1 (1979 1-108. iv 22 Géographie Botanique, with an English translation quickly following. This book listed at least twenty-one species of plants and algae he believed were carried between lands or seas by human agency. One ongoing debate of the time concerned the number of different “centers of creation” needed to reconcile the Genesis creation account with life’s geography and variety. While declaring “All testimonies, indeed, confirm the suspicion that the human race, and the domestic animals, descended from the high mountain plains of Central Asia” (tacitly, Eden), de Candolle dismissed the idea that all plant species had radiated likewise from a single Asian source as “physically impossible.”31 He concluded “the vegetable world has neither descended from one common birthplace, nor diffused itself from one country into another; but that vegetation is in every case the product of the joint influence of temperature, soil, and the particular composition of the moisture of the earth.”32 Charles Lyell was impressed with the effort. “Decandolle has enumerated twenty great botanical provinces inhabited by indigenous or aboriginal plants; and although many of these contain a variety of species which are common to several others, and sometimes to places very remote, yet the lines of demarcation are, upon the whole, astonishingly well defined.” 33 The idea that plants (and animals) somehow belong to places predated de Candolle and 31 Augustin P. de Candolle, Elements of the Philosophy of Plants, in Lomolino, et al, eds. Foundations of Biogeography, (see cit. n. 26) 45. 32 Ibid., 48 33 Lyell, Principles of Geology, (cit. n. 22) 71 iv 23 remains current, even though the justifications for this belief have evolved with the times. Charles Lyell Charles Lyell considered himself a geologist, but was (as suggested above) a keen follower of debates regarding the origin and distribution of life. This “greater geology” led him to comment, usually astutely, on other branches of natural history, and his comments help to contextualize the pre-Darwinian debates. First, with regard to the problems of geography and creation: The first travellers were persuaded that they should find, in distant regions, the plants of their own country, and they took a pleasure in giving them the same names. It was some time before this illusion was dissipated; but so fully sensible did botanists at last become of the extreme smallness of the number of phaenogamous plants common to different continents, that the ancient Floras fell into disrepute. All grew diffident of the pretended identifications, and now we find that every naturalist is inclined to examine each supposed exception with scrupulous severity. [N: de Candolle, Essai Elém. de Géog. Botan. p. 45.] If they admit the fact, they begin to speculate on the mode whereby the seeds may have been transported from one country into the other, or inquire on which of the two continents the plant was indigenous, assuming that a species, like an individual, cannot have two birth-places.34 Lyell referred to neobiota several times. After discussing various modes of plant dispersal: “In addition to all the agents already enumerated as instrumental in diffusing plants over the globe we have still to consider man– one of the most important of all. He transports with him, into every region, the vegetables which he cultivates for his wants, and is the involuntary means of spreading a still greater number which are useless to him, or even noxious.”35 34 Ibid. 35 Ibid., 82 iv 24 Slightly later, “The most remarkable proof, says de Candolle, of the extent to which man is unconsciously the instrument of dispersing and naturalizing species, is found in the fact, that in New Holland, America, and the Cape of Good Hope, the aboriginal European species exceed in number all the others which have come from any distant regions, so that, in this instance, the influence of man has surpassed that of all other causes which tend to disseminate plants to remote districts.”36 Taking a stance both pragmatic and ethical, Lyell related neobiota and species extirpations in a proto-conservationist statement: If we drive many birds of passage from different countries, we are probably required to fulfill their office of carrying seeds, eggs of fish, insects, molluscs, and other creatures, to distant regions; if we destroy quadrupeds, we must replace them, not merely as consumers of the animal and vegetable substances which they devoured, but as disseminators of plants, and of the inferior classes of the animal kingdom. We do not mean to insinuate that the same changes which man brings about, would have taken place by means of the agency of other species, but merely that he supersedes a certain number of agents, and so far as he disperses plants unintentionally, or against his will, his intervention is strictly analogous to that of the species so extirpated.37 On the other hand, Lyell saw neobiota — and their human dispersers — as essentially natural. Drawing on Robert Burns’ poem “To a Mouse, on Turning Her Up in Her Nest with a Plough” and Shakespeare’s As You Like It, he stated his case: If we wield the sword of extermination as we advance, we have no reason to repine at the havoc committed, nor to fancy, with the Scotch poet, that 'we violate the social union of nature;' or complain, with the melancholy Jaques, that we 36 Ibid., 84 37 Ibid. iv 25 'Are mere usurpers, tyrants, and, what's worse, To fright the animals, and to kill them up In their assign'd and native dwelling place’ We have only to reflect, that in thus obtaining possession of the earth by conquest, and defending our acquisitions by force, we exercise no exclusive prerogative. Every species which has spread itself from a small point over a wide area, must, in like manner, have marked its progress by the diminution, or the entire extirpation, of some other, and must maintain its ground by a successful struggle against the encroachments of other plants and animals. That minute parasitic plant, called 'the rust' in wheat, has, like the Hessian fly, the locust, and the aphis, caused famines ere now amongst the 'lords of the creation'. The most insignificant and diminutive species, whether in the animal or vegetable kingdom, have each slaughtered their thousands, as they disseminated themselves over the globe, as well as the lion, when it first spread itself over the tropical regions of Africa.38 Lyell summarized his attitude by putting humans in context, in what may yet emerge as the most astute possible conclusion regarding the biogeographic influence of the species: “we may regard the involuntary agency of man as strictly analagous to that of the inferior animals. Like them we unconsciously contribute to extend or limit the geographical range and numbers of certain species, in obedience to general rules in the economy of nature, which are for the most part beyond our control.”39 Anekeitaxonomy: A Tradition Emerges Some botanist-phytogeographers applied themselves to “adventive floristics,” categorizing plants according to degrees of establishment, modes of dispersal, and even by the intentions of those who transported them.40 I have 38 Ibid., 156 39 Ibid., 122 40 Herbert Sukopp, “On the Early History of Urban Ecology in Europe,” Preslia, Praha 74 (2002): 373-393. iv dubbed the resulting tradition “anekeitaxonomy.” 41 26 Grounded in the phytogeographical speculations and observations of Buffon, Willdenow, Humboldt, and (especially) A.P. Candolle, it was founded in earnest by the irascible Englishman Hewett Cottrell (H.C.) Watson. Candolle’s son Alphonse independently developed its ideas further. A later Swiss botanist, Albert Thellung, standardized the terminology and took it into the twentieth century; some have commemorated the resulting system as the “Thellungian Paradigm.” Anekeitaxonomy persisted in Western Europe in (e.g.,) the work of Morten Porsild in Greenland, and in the United States in Victor Mülenbach’s botanical surveys of Missouri railroad yards.42 Some of its terminology persists in modern usage, particularly in Central Europe and Russia. Petr Pyšek of the Department of Invasion Ecology at the Institute of Botany, Academy of Sciences of the Czech Republic has most recently shouldered the burden of preserving the anekeitaxonomic tradition, and his account of its purposes and history is usefully laconic and frank: More than a century of effort put into the classification of manaccompanying plants by European botanists has yielded a number of classification systems ([Alphonse] de Candolle, 1855; Ascherson 1883; Rikli 1903; Thellung 1922; Schroeder 1969; Holub and Jirásek 1967; 41 Categorizing biota by proposed geographical attachment; from ανήκει, “belonging”, in the spirit of their often Grecianized terminology.) There is also an obscure, opportunistic pun at play here. Michael Zohary, perhaps the last of the unselfconscious anekeitaxonomists, closed out the era with a shrug by employing the term “anecophytes” (presumably from αν-οικος, or home-less) for “plants whose original habitat is unknown.”i Plant Life of Palestine, Israel and Jordan (New York: Ronald Press 1962): 219. 42 Morten P. Porsild, Alien Plants and Apophytes of Greenland. (Copenhagen: C.A. Reitzels, 1932); Mühlenbach, "Contributions to the Synanthropic (Adventive) Flora of the Railroads in St. Louis, Missouri, U.S.A." (cit. n. 30). iv 27 Kornas 1990). These are mostly based on (1) time of immigration of a species into the region, (2) the means of introduction by humans, i.e., intentionally or unintentionally, and (3) degree of naturalization, i.e., its ability to become established under local conditions … Some of these systems are rather complicated and have led to the creation of an extensive jargon … Their complexity is probably the main reason they did not come into wider use. Some of them (e.g., Kornas 1990), by defining a number of categories often based on vague criteria, impose a number of practical problems that emerge if one is trying to use them, e.g., how to distinguish seminatural and natural habitats or when is an alien already permanently established and when not.”43 Anekeitaxonomy, following the tradition of A.P. de Candolle and many others, assumes that plant species initially occurred in, and effectively belonged to a particular geography, and humans moved some species to new locations. This concept was well established (e.g., Willdenow’s mountaintop floral centers) before the publication of On the Origin of Species and somehow robust enough to survive the Darwinian revolution without much difficulty. Hewett Cottrell (H.C.) Watson H.C. Watson’s early, primary interest was phrenology, and he was frustrated by the general failure of that study. But he was an accomplished botanist, and by most accounts, an equally accomplished misanthrope.44 A regular correspondent and sounding-board for Charles Darwin in the period leading up to publication of the Origin of Species, he was concerned that natural selection was held to account for more than it actually accomplished.45 Among 43 Petr Pyšek. “On the Terminology used in Plant Invasion studies,” IN Peter Pyšek, et al, Eds. Plant Invasions, General Aspects and Special Problems (Amsterdam: SPB Academic, 1995) 72-73. 44 See Frank N. Egerton, Hewett Cottrell Watson: Victorian Plant Ecologist and Evolutionist. (Aldershot: Ashgate, 2003). 45 Ibid., 177-193 iv 28 his own accomplishments, Watson established the first relativistic “formula” or taxonomy of phytogeographic classification.46 Established, rather than coined, because after codifying it in the first volume of his Cybele Britannica, he applied and refined it extensively, as did some of his botanic successors. Relativistic, because his labels concern the origins of plant species vis-à-vis their presence on the island of Britain (the contiguous regions of England, Scotland and Wales), rather than assigning species of extra-British origin to particular realms. By 1847 botanists were recognizing that plant species had lately appeared on the island, but they lacked terms to distinguish various degrees of establishment. The term native was traditionally used to describe untamed, uncultivated spontaneously growing vegetation, regardless of particular species or suspected origins. But Watson, working specifically on the distribution of plants in Britain, was dissatisfied with both the accuracy and precision of knowledge of these matters. “Among the plants now found seemingly wild in Britain, whether more or less widely distributed, are many species which occur under conditions calculated to suggest an idea that they may have been first introduced into this island by the agency of mankind.”47 He considered the indications of origin found in general British Floras to be “little better than idle 46 H.C. Watson, Compendium of the Cybele Britannica. Part First (Thames Ditton: privately published 1868) 62; Watson employed the term phytogeography. 47 Ibid., 59. iv 29 guesses, usually hazarded without any sufficient qualifications for judgment in their authors.”48 In Watson’s view, competition among “vainglorious” botanizers to locate and identify new British species and new occurrences of known ones was increasing the confusion. His solution was a series of classifying terms describing the “civil claims and local situation” of every species found on the island.49 Some of his terms are seemingly self-evident, and persist in modern usage; quite often, perhaps, without reference to or awareness of Watson’s definitions. Others, more idiosyncratic, never gained much currency; even Watson himself dropped two of them and added another during the first twenty years of their application. Watson’s formula follows. Refinements or reformulations from the 1868 version are italicized; minor alternative wordings appear in brackets {}, as do 1847 words dropped in 1868. Species names exemplifying each class are omitted here. 1. Native: Apparently an aboriginal British species; there being little or no reason for supposing it to have been introduced by human agency. 2. Denizen: At present maintaining its habitats, as if a native, without the aid of man, yet liable to some suspicion of having been originally introduced by human agency, whether by design or by accident. 3. Colonist: A weed of cultivated land, by road sides or about houses, and seldom found except in places where the ground has been adapted for its production and continuance by the operations of man; with {a / some} tendency, {however, /also in some of them} to appear {also} on the shores, landslips, {&c.} and what are called “waste places. [five species] are weeds of cultivated land, and would perhaps disappear if plough and spade ceased their work. Several [species of one genus, plus four of 48 H.C. Watson, Cybele Britannica Vol. IV (London: Longman & Co, 1859) 70. 49 Ibid.; H.C. Watson, Cybele Britannica Vol. I (London: Longman & Co. 1847) 63 iv 30 others] constitute connecting links between the Colonists and Denizens, found chiefly by road sides, rubbish heaps dunghills, and near the sea. 4. Alien: Now more or less established, but either presumed or certainly know to have been originally introduced from other countries. 4. Alien species are those certainly or very probably of foreign origin; though several placed in this category are now well established amid the indigenous flora of this island; others less perfectly so. 5. Incognita: Reported as British, but requiring confirmation as such. Some of these have been reported through mistakes of the species…Others may have been really seen in the character of temporary stragglers from gardens…Others cannot now be found in the localities published for them…though it is not improbable that some of these may yet be found again. A few may have existed for a time, and become extinct. 5. Casual species are chance stragglers from cultivation; those occasionally imported and sown with agricultural seeds; those introduced among wool, oil-seeds, or other merchandize [sic]; foreign plants found on ballast heaps deposited from ships; and generally such alien species as are most uncertain in place or persistence. 6. Hibernian, or Sarnian: Native, or apparently so, in Ireland, or in the Channel Isles, though not found in Britain proper. 6.[Not used.] Neither Watson’s biographer Frank Egerton nor I found reason to suspect that this formula was essentially chauvinistic. To our knowledge, Watson never stated or suggested that his British natives were inherently superior to the others.50 However, by describing his formula in part as one of “civil claims,” and elsewhere as “predial” (i.e., landholding) Watson did seem to regard inclusion in the native British flora in combined terms of occupancy rights and a kind of botanical citizenship.51 He emphasized, “the distinction between native and introduced species is absolute and real,” but seemingly did not extend the 50 Frank Egerton to Matt Chew, 27 September 2005. 51 Watson, Cybele I, (cit. n. 49) 63; Cybele IV, (cit. n. 48) 107. iv 31 argument to propose defending native British plants against more recent comers on the basis of prior or otherwise superior claims.52 When the fourth volume of his Cybele Britannica appeared, Watson had, or believed he had, several rival formulations to contend with. Chief among his list of pretenders were Alphonse de Candolle [see below], and the Hookers, W.J. (father) and J.D. (son) along with their collaborators. Watson criticized de Candolle at some length for what might best be summarized as an incautious reliance on dubious sources; but praised him in specific instances.53 Watson also commented on de Candolle’s decision to collect opinions rather than synthesize them according to a rigidly defined formula. Introducing an extended, species-by-species table comparing his own judgments with de Candolle’s, Watson wrote, “the third column shows the opinion of M. De Candolle compressed into one or two words, so far as this can be done from the remarks made on the plants in the Geographie Botanique.”54 By 1859 Watson clearly felt that the point of his foray into “civil claims’ was to purge the list of British plants of species he considered alien according to his own definition. He closed volume IV, chapter III of Cybele Britannica with a paragraph that suggests much about his interests, opinions and character: This subject of introduced plants has encroached too largely on the pages of the present work, and must now be left for other topics. The Author has hitherto unsuccessfully tried to induce some other botanist to write a ‘History of the British flora’;— tracing out each species back to the 52 Ibid., 68. 53 Ibid., 83 ff. 54 Ibid., 97. iv 32 earliest records of its occurrence in Britain; and also, when possible, to its still earlier relics in peat-mosses and elsewhere. The vast changes gradually wrought in the vegetation of Britain, by the conversion of forests into wastes, and of wastes into cultivated lands, would also find appropriate place in a work of the kind suggested. Such a history might be made a very valuable contribution to botanical science,— and likely enough to geological science also. But it would require to be written by a scrupulous investigator of facts, and a conscientious recorder of them. It would require to be written under a higher impulse than that which prompts to the compiling of ‘Keys’ and ‘Manuals’ for the market,— or under a worthier ambition than that of “getting up some sort of paper, likely to make a talk among the geologists, at the next meeting of the British Association.”55 Eventually Watson settled on a distinction that still underpins invasion biology. He differentiated between two basic approaches to the study of flora, and threw in his lot with the latter; answering the two, trailing “how” questions would consume his successors. 1. Geographical Botany is understood to begin with the plants themselves, whether by individual species or in generic or ordinal groups, and to trace the distribution of the species or groups over the surface of the Earth, or over any portion of it immediately under consideration; 2. Botanical Geography [floristics] regards the Earth’s surface itself primarily, and examines the floral peculiarities of its various parts or divisions; investigating the diversities and correspondences of their respective floras, and endeavouring to ascertain the circumstances or influences which have determined the existing conditions of plant-distribution. Briefly, the former may be said to treat about the places of the plants; the second, conversely, to treat about the plants of the places. If reduced into brief questions, the distinction may become still more clear. 1. In what places, and under what conditions, does this plant or group of plants occur? 2. By what kind of plants, or combination of plants, is this country inhabited and characterized? — how did they first get to it? — how maintained there?” 56 55 Ibid., 124-5. 56 Watson, Compendium, (cit. n. 46), 1. iv 33 Alphonse de Candolle A. P. de Candolle’s son Alphonse continued and expanded his father’s work, and succeeded him as Director of the Geneva Botanical Garden. He, too, is relatively well known and will not be introduced at length. H.C. Watson considered anyone with a different point of view to be an antagonist, and I suggested this by including his mention of de Candolle, above.57 In 1855, as part of a massive, two-volume work, de Candolle published an alternative to Watson’s system. He explicitly compared it to Watson’s categorization, but not in a particularly vehement manner. From [Alphonse de Candolle’s] point of view, the species that live in a country subdivide in this manner: They are firstly either cultivated or spontaneous. The CULTIVATED species are of two kinds: 1. Cultivated voluntarily 2. Cultivated involuntarily (bad grasses [weeds], unquestionably parasitic). I speak here about the species which absolutely exist only in the fields, gardens, etc., without being in open country in a spontaneous state, because one cultivates also indigenous plants, and there are usually spontaneous species which are seen sometimes in the cultivated grounds of their own country. The SPONTANEOUS species, i.e., which live and propagate without the help of the man, belong to various categories, which I will enumerate, while starting with the least important and those which have the strongest relationship with the crop plants. These plants are: 1. Adventitious, i.e., foreign of origin, but badly established, being able to disappear from one year to another. 2. Certainly naturalized, and by abbreviation, naturalized. They are well established in the country, but there is positive evidence of a foreign origin. 3. Probably of foreign origin. Well established species; but according to strong indications, there are more reasons to believe them of origin foreign 57 Egerton, Hewett Cottrell Watson, (cit. n. 44) 1-4. iv 34 than primitive in the country. In other words, to restate my idea more clearly, the odds favoring a foreign origin are better than even. 4. Perhaps of foreign origin. There are in such cases some indications of a foreign origin, though the species are long and well established in the country. For one reason or another, one can raise some doubts about their indigenousness. 5. Indigenes, aboriginals, native (simultaneously an English and French expression). All the spontaneous species whose origins are not doubtful, which appear to have existed in the country before to the influence of man, probably for geological rather than historical time; for example, if in Great Britain, since before the separation of this island from the continent.58 Later in Geographie, de Candolle made an interesting summary remark about the human disperal of plants: “Before the appearance of man, before the multiplicity of his effects, naturalizations were infinitely rare, because they are this very day in Europe, and when they arrive, it is almost always by our direct or indirect influence.”59 This uniformitarian approach suggests Lyell’s influence. Alphonse de Candolle’s distinction between cultivated and uncultivated plants in Géographie presaged a later major work. Mid-nineteenth century phytogeographers were absorbed, if not obsessed, with discovering the origins of cultivated plants. Candolle attacked the problem in substantial fashion with The Origin of Cultivated Plants. This study simply assumed human agency as a primary factor, but Candolle made several explicit comments: “Naturalized plants spread rapidly. I have quoted examples elsewhere … of instances within the last two centuries, and similar facts have been noted from year to year. The rapidity of the recent invasion of [a species] into the rivers of Europe is well known, and 58 Alphonse de Candolle, Geographie Botanique Raisonée (Paris: V. Masson, 1855) Vol. 2., 642-643. My translation. 59 Ibid., 709. iv 35 that of many European plants in New Zealand, Australia, California, etc. mentioned in several floras or modern travels.”60 He also proposed a dichotomy between “normal” rates of increase for new arrivals and decrease for failing types: “The great abundance of a species is no proof of its antiquity … As a rule, an invading species makes rapid way, while extinction is, on the contrary, the result of the strife of several centuries against unfavourable circumstances.”61 Joseph Dalton Hooker Discussions of H.C. Watson and Alphonse de Candolle inevitably include J.D. Hooker as a significant interlocutor. Prefacing the second edition of his Student’s Flora of the British Islands, Hooker deferred, more or less, to an earlier formula: “To the doubtfully indigenous Species I have often added Watson’s opinion as to whether they are ‘aliens,’ ‘colonists,’ or ‘denizens,’ &c. It may be well to repeat here his definitions of these terms, premising that by ‘native’ is meant that the Species has not been introduced by human agency:— A denizen is a Species suspected to have been introduced by man, and which maintains its habitat. A colonist is one found only in ground adapted by man for its growth and continuous maintenance. An alien has presumably been introduced by human agency.”62 Hooker also added a footnote, “The vagueness of these definitions is unavoidable; and their correct application in many cases is exceedingly difficult. Few who have not gone into the subject have an idea of how many plants would disappear from our Flora were the soil left undisturbed by man and the lower 60 Alphonse de Candolle, Origin of Cultivated Plants (New York: D. Appleton, 1885), 13. 61 Ibid., 13 62 Joseph D. Hooker, A Student’s Flora of the British Islands (London: MacMillan, 1884), vii. iv 36 animals which he rears. I think it probable that both the Shepherd’s-Purse and the common form of the Dandelion would be amongst the first to be suppressed.”63 Documentation so far acquired suggests that Hooker had ideas about anekeitaxonomy, but never devised categories of his own. His 1866 lecture on “insular floras” included several potential categories, but none was explicitly defined. Hooker used the terms “native” and “indigenous” synonymously, and in the typical sense; at one point he mentioned “bonâ fide natives,” and at another, “truly indigenous” when discussing the work of other botanists. Hooker’s use of “cultivated,” “semi-naturalised,” “naturalised” and “quasi-indigenous” might have represented a continuum. There he applied “introduced” only as a verb; “exotic” seems to have been his adjective of choice at that point for introduced plants. An appendix to the Student’s Flora listed “excluded species, applying many terms in addition to (or as synonymous with) the Watsonian system. Hooker apparently coined the term “escape” (as both noun and verb) for cultivated plants that spread beyond their intended locations.64 Among the more and less definite adjectives from the appendix: “only where planted,” “reported,” “found,” “introduced,” “a cottage ornament,” “a garden relic,” “almost naturalized,” “naturalized,” and the very specific yet somehow still vague “shrubberies” or “only in shrubberies.”65 63 Ibid., vii, footnote. 64 Oxford English Dictionary Online 65 Hooker, A Student’s Flora, (cit. n. 62) 527-535. iv 37 Hooker was notable, for his period, for his evident dismay over the conversion of island floras (specifically: Ascension, the Azores, Canaries, the Cape Verdes, Desolation, the Madeiras, and St. Helena) by introduced domesticated herbivores (particularly goats) and European plants. His concern seemed to be mainly over the loss to science, but his rhetoric was inflammatory, and seemed meant to incite an emotional response. Hooker’s account of St. Helena was particularly vehement, anticipating in tone those of much later writers; but also echoing in many respects a then-recent description by an American George Perkins Marsh.66 The Spanish introduced goats to St. Helena early in the 16th century. Hooker gave an extended account of the aftermath, from which this is extracted: [Goats] were introduced in 1513, and multiplied so rapidly, that in 1588 … they existed in thousands; single flocks being almost a mile long … in 1709, trees still abounded … however, the Governor of the island reported … that the timber was rapidly disappearing, and that the goats should be destroyed … He received the laconic reply "The goats are not to be destroyed, being more valuable than [trees]" … in 1810 another Governor reports the total destruction of the great forests by the goats … still, even then, so great was the amount of seed annually shed, so rich the soil, and so rapid the growth of the native plants … that if the goats were killed, and the island left to itself, it would in 20 years be again covered with indigenous vegetation … the goats were killed, but another enemy to the indigenous vegetation was at the same time introduced, and which has now rendered it in all probability impossible that the native plants will ever again resume their sway. [The] Governor … proposed and carried cut the introduction of exotic plants on a large scale, and from all parts of the world; these have propagated themselves with such rapidity, and grown with such vigour, that the native plants cannot compete with them. The struggle for existence had no sooner begun, than the issue was 66 Marsh was not a scientist in the sense assumed for purposes of this paper. If its scope were expanded to include Marsh, it would also include R.W. Emerson, Victor Hehn, Ben-Hur Lampman, John Muir, Charles Pickering, H.D. Thoreau, H.G. Wells, and many others. iv 38 pronounced; English Broom, Brambles, Willows and Poplars Scotch Pines and Gorse bushes, Cape of Good Hope bushes, Australian trees and American weeds, speedily overran the place; and wherever established, they have actually extinguished the indigenous Flora … it is therefore now impossible to distinguish the introduced from the native plants of St. Helena … probably 100 St. Helena plants have thus disappeared from the Systema Naturæ since the first introduction of goats on the island. Every one of those was a link in the chain of created beings, which contained within itself evidence of the affinities of other species, both living and extinct, but which evidence is now irrecoverably lost.67 George Bentham Since today’s botanists seem to have forgotten them, it is tempting to assume that neobiota were of marginal interest to their nineteenth century precursors. But there is evidence to the contrary. While compiling his “popularlevel” Handbook of the British Flora, George Bentham (nephew of the betterknown Jeremy) wrote a twenty-seven page, essay review of de Candolle’s Géographie.68 Much of it was commentary on attempts to reconcile the increasing weight of empirical evidence with traditional, Biblical creation. For Bentham, like de Candolle and many others, the recent history of plant geography was in many ways an entrée to discussing the history and origin of life on earth, a topic of intense debate at the time. Bentham made it clear that explaining geographical distribution was a significant concern and frustrating bone of contention among botanists: “numerous writers of local floras, special monographs and other partial botanical 67 Joseph D. Hooker, “Insular Floras” Gardeners' Chronicle January 1867, accessed online at http://www.wku.edu/~smithch/biogeog/HOOK1866.htm, 12 October 2006. 68 P. F. Stevens, “Bentham, George (1800–1884).” Oxford Dictionary of National Biography. H. C. G. Matthew and Brian Harrison, Eds. Oxford: University Press, 2004. iv 39 works or memoirs, whilst they have each contributed invaluable data towards the general consideration of the subject, have, at the same time, been but too frequently tempted to generalise hastily their own partial observations. The result has been to supply us with endless speculations on botanical regions, on endemic and sporadic species, on their development and migrations, with a formidable array of figures and tables, founded upon incomplete and deceptive data, which have tended to bewilder rather than enlighten us on the true principles of the science.”69 However, that lamentable state of affairs did not prevent Bentham from offering views of his own. These, incidentally, are hardly distinguishable from many put forth 150 years later: In our own days … physical obstacles are occasionally overcome by the direct or indirect agency of man, Immense tracts of land are devoted by him expressly to the cultivation of plants which did not previously exist there. In cultivating these, he has unintentionally sown others which come up as weeds; seeds borne by these cultivated plants, or accidentally carried by man with his goods, &c., in the course of his migrations, have sown themselves and established their species in any stations [i.e., habitats] suited to their constitutions where they may have been dropped. In some few instances we may observe, even now, the area of species extended by causes purely natural; but in the great majority of instances, where man has not exercised his influence, the area of species appears to have remained the same for a period going far beyond our historical times, and commencing probably long previous to the creation of man.70 Bentham went on, offering his own revision of de Candolle’s categorization: “Hence the division of the plants of a district into cultivated plants, either by the express will of man, or as weeds of cultivation, ‘cultivated in spite of the will of man:’ and wild plants, which are termed naturalised if they have 69 George Bentham, “de Candolle’s ‘Geographical Botany’” Edinburgh Review or Critical Journal 104 (1856) 490-518 70 Ibid. iv 40 established themselves after their introduction by the agency of man direct or indirect, indigenous if we believe them to have existed in the country previous to the operation of human agency.”71 In general, later anekeitaxonomists built on both the Watson and de Candolle conceptions, and Bentham’s version was probably typical of many others. Like de Candolle they desired to account for and classify domesticated species; like Watson they preferred positively defined categories like alien to the more negatively defined “doubtful” ones of Candolle. They also echoed Watson’s borrowings of “civil” status. Perhaps this led its practitioners to desire ever more specific detail, and the greater exclusion that resulted required the corresponding creation of new positives. Continuing the regression would inevitably tend to generate a special category for each species; an unwieldy model, indeed. Albert Thellung and His Successors Albert Thellung was another Swiss systematist-phytogeographer, educated and working within the “Zurich-Montpellier school” of botany. In addition to taxonomic accounts, he contributed to a major regional flora (Illustrierte Flora von Mittel-Europe, 1906) and then developed a monograph on the many adventive species growing around a major wool-processing center, Port Juvenal (the harbor of Montpellier, France). Using dated inventories, he was able to correlate the commercial history of the port with the progression of 71 Ibid. iv 72 foreign plants present in the area. 41 He also assisted Ida Hayward and George C. Druce with specimen identifications for another, 1919 “wool garden” inventory conducted along the Tweed and Gala rivers in southwestern Scotland, garnering that book’s dedication for his “kindly help.”73 Thellung died suddenly and relatively young in 1928.74 The Zurich-Montpellier school of botany was afterward dominated by the “plant sociology” of Josias Braun-Blanquet, who edited Thellung’s last major publication, another in the still-ongoing tradition of speculating on the origins and emergence of cultivated plants.75 As Pyšek noted, anekeitaxonomies have become complex; much of that complexity resides in a terminology that expands with every identified (or assigned) permutation of immigration date, means (and intentionality) of introduction by humans, and degree of naturalization. A quick literature review identified over 250 extant compound concepts, labeled with Greek (or pseudoGreek) neologisms.76 It is not yet entirely clear who initiated the practice of Greek terminology for these purposes, but the idea was consistent with the familiar classicizing tradition of taxonomic nomenclature. Citations in Thellung’s Adventive Flora of Montpellier suggest that it can be traced back at least to 72 Mühlenbach, "Contributions” (cit. n. 30). 73 Ida M. Hayward and George C. Druce The Adventive Flora of Tweedside (Arbroath, Scotland: T. Buncle 1919). 74 Hans Schinz "Albert Thellung 1881-1928." Vierteljahresschrift der Naturforschenden Gesellschaft in Zürich 73 (1928): 558-80. 75 Albert Thellung, Die Entstehung der Kulturpflanzen (Munich: F.P. Datterer, 1930). 76 These are considered in greater detail in M. Chew, “Anekeitaxonomy: Biologizing Belonging” in prep. iv 77 German botanist Paul Ascherson in 1883. 42 Ascherson, like Watson, had compiled a regional flora, suggesting that the need to account for casual additions was a recurring feature of arbitrarily bounded inventories.78 Albert Thellung’s 1912 descriptions of three of anekeitaxonomy’s most basic categories (anthropochores, neophytes, and epœkophytes) are translated here, because more recent published summaries have discarded some of the original system’s more revealing characteristics. Constituting perhaps one percent of the present system’s named concepts, they nevertheless exemplify its problematic complexity. “Anthropochores” are “Plants which are propagated while benefiting from the action of man. This division includes all the species which are not originally indigenes (indigenous) in the area in question, but which were introduced there by man, intentionally or without his knowledge; they are thus the cultivated species and the bad grasses [i.e., weeds] of foreign origin.”79 “Neophytes,” a subclass of anthropochores, are “new citizens, having acquired middle-class [Neubürger] status. Plants reproducing more or less abundantly and constantly in natural sites, forming part, seemingly, of the indigenous vegetation; their future existence is thus independent of man.”80 77 Paul Ascherson in Johannes Leunis Synopsis der Pflanzenkunde 3, I (1883) 791-796; cited in Albert Thellung, La flore adventice de Montpellier (Cherbourg, Le Maout, 1912). 624 n.7. 78 See Paul Ascherson, Flora der Provinz Brandenburg, der Altmark und des Herzogthums Magdeburg (Berlin: A. Hirschwald, 1864). 79 Thellung, La flore adventice, (cit. n. 77) 625-628; My translation. 80 Ibid. iv 43 Epœkophytes, another anthropochore subclass, are “colonists [Ansiedler, colloquially, ‘squatters’]. Species present since recent times and also constantly reproducing, but restricted to artificial localities (e.g., especially liking fields and vineyards). The persistence of these species depends on man, who must preserve (or rather continuously re-create) conditions appropriate to them (plowed ground, cleared patches); they produce ripe seeds in sufficient quantity, but they cannot compete with invading indigenous species.”81 Fraught with multiple causation and incommensurable criteria, anekeitaxonomies are not wholly hierarchical. They include a wide range of difficult-to-accommodate categories, from (e.g.,) polemechores (plants spread by warfare) to chomophytes (fissure or ledge plants). But it is worth noting that even though this system has aspects as artificial as those of classic Linnaean taxonomy, when the two are combined they read rather like a natural method of classification. Anekeitaxonomy can thus be seen as one kind of attempt to move general plant taxonomy in a more intuitive and practical direction. Other Phytogeographical Accounts Charles Darwin’s Beagle voyage journal entry for 19 September 1833 provided the earliest application I have seen of the term invasion to a population of introduced plants. It specifically referred to cardoon (i.e., wild artichoke) plants, but opened a general reflection on the effects of what Hooker would later call “escaped” (i.e., feral) domesticated plants and animals. The only other generalized terms Darwin used in the immediate context were “aborigines” and 81 Ibid. iv 44 “introduced,” so it did not rise to the level of anekeitaxonomy. Nevertheless, it was Darwinian, and that lends interest to an otherwise relatively mundane set of observations. Near the Guardia we find the southern limit of two European plants, now become extraordinarily common. The fennel in great profusion covers the ditch-banks in the neighbourhood of Buenos Ayres, Monte Video, and other towns. But the cardoon (Cynara cardunculus) has a far wider range: it occurs in these latitudes on both sides of the, Cordillera, across the continent. I saw it in unfrequented spots in Chile, Entre Rios, and Banda Oriental. In the latter country alone, very many (probably several hundred) square miles are covered by one mass of these prickly plants, and are impenetrable by man or beast. Over the undulating plains, where these great beds occur, nothing else can now live. Before their introduction, however, the surface must have supported, as in other parts, a rank herbage. I doubt whether any case is on record of an invasion on so grand a scale of one plant over the aborigines. As I have already said, I nowhere saw the cardoon south of the Salado; but it is probable that in proportion as that country becomes inhabited, the cardoon will extend its limits. The case is different with the giant thistle (with variegated leaves) of the Pampas, for I met with it in the valley of the Sauce. According to the principles so well laid down by Mr. Lyell, few countries have undergone more remarkable changes, since the year 1535, when the first colonist of La Plata landed with seventy-two horses. The countless herds of horses, cattle, and sheep, not only have altered the whole aspect of the vegetation, but they have almost banished the guanaco, deer and ostrich. Numberless other changes must likewise have taken place; the wild pig in some parts probably replaces the peccari; packs of wild dogs may be heard howling on the wooded banks of the less-frequented streams; and the common cat, altered into a large and fierce animal, inhabits rocky hills. As M. d'Orbigny has remarked, the increase in numbers of the carrionvulture, since the introduction of the domestic animals, must have been infinitely great; and we have given reasons for believing that they have extended their southern range. No doubt many plants, besides the cardoon and fennel, are naturalized; thus the islands near the mouth of the Parana, are thickly clothed with peach and orange trees, springing from seeds carried there by the waters of the river.82 82 Charles Darwin. The Voyage of the Beagle (Washington DC: National Geographic Society, 2004.) 104-105 iv 83 Darwin’s next recorded thoughts dealt with actual military matters. 45 It is interesting to speculate whether the militarized milieux of his childhood during the later Napoleonic Wars, then-recent wars of South American independence, and ongoing military actions against hostile indigenous peoples during his visit influenced Darwin to see and describe cardoon as an invading force. Consider, too his allusion to this episode in the Origin: Cases could be given of introduced plants which have become common throughout whole islands in a period of less than ten years, Several of the plants now most numerous over the wide plains of La Plata, clothing square leagues of surface almost to the exclusion of all other plants, have been introduced from Europe; and there are plants which now range in India, as I hear from Dr Falconer, from Cape Comorin to the Himalaya, which have been imported from America since its discovery. In such cases, and endless instances could be given, no one supposes that the fertility of these animals or plants has been suddenly and temporarily increased in any sensible degree. The obvious explanation is that the conditions of life have been very favourable, and that there has consequently been less destruction of the old and young, and that nearly all the young have been enabled to breed. In such cases the geometrical ratio of increase, the result of which never fails to be surprising, simply explains the extraordinarily rapid increase and wide diffusion of naturalised productions in their new homes.84 Here Darwin’s account dwelled on the favorability of circumstances affecting the introduced species rather than calling it an “invasion” and subjugation of the “aborigines,” although later in the Origin he suggested that these European plants had “to a certain extent beaten the natives.”85 Since his purpose was to provide examples of selection, these and other brief mentions of neobiota in the 83 Ibid., 105-106 84 Charles Darwin, On the Origin of Species by Means of Natural Selection (London: John Murray, 1859), 63-64. The sixth edition mentioned cardoon specifically, but the differences in the versions are too slight to concern us here. 85 Ibid., 379-380. iv 46 Origin tended not to dwell on deploring the process under study. It is also, perhaps notable that Darwin framed the Origin wholly during the so-called Pax Britannica when British culture was less warfare-oriented during this period than at practically any time previously. If it had happened otherwise, perhaps Darwin’s “struggle” for existence would have been framed as a “battle.” Douglas H. Campbell Douglas Campbell was an original (1891) member of the Stanford University faculty and longtime chair of the Botany Department. His specialty was bryology, (the study of mosses) and his contributions to that field were well respected.86 In 1926 he forayed into the wider world with An Outline of Plant Geography. The book received an apologetic but exquisite panning from M.L. Fernald in Ecology (to which the journal’s editor appended a concurring footnote).87 Further evidence of the book’s poor reception was its omission from Campbell’s list of published works in his obituary.88 Campbell’s phytogeographical acumen is not at issue here. In factual terms, he merely summarized much that had been said (sometimes long) before, without improving on it. But he is significant to my thesis because he appears to have been one of science’s early adopters of a misanthropic, nativist attitude 86 William C. Steere, “Douglas Houghton Campbell (1858-1953)” The Bryologist 56 (1953) 127-133. 87 M.L. Fernald, “Campbell’s Outline of Plant Geography” Ecology 7(1926) 4:510516. Fernald’s closing thought: “By its publication the day when phytogeography shall become a science rather than a pleasant pastime is still further postponed.” 88 Steere, “Douglas Houghton Campbell (1858-1953)” (cit. n. 86). iv 89 toward the neobiotic plants. 47 Several quotations demonstrate this, in both style and substance. After listing various modes of dispersal, Campbell wrote, “Finally, man, involuntarily or otherwise, has been the cause of the migration of many plants over pretty much the whole earth.”90 He then elaborated with a ‘fall of man’ assessment: “In his earliest development primitive man only took from the forest such food as he might find–fruits, nuts or roots, thus doing no more to disturb the equilibrium of the plant association than would be done by the foraging of any other animal; but as soon as he developed the most rudimentary form of agriculture, or attempted to provide grazing for animals, tame or wild, he began a warfare with the vegetable kingdom which has continued with increasing energy down to the present time.”91 Like many who would follow, he offered a postfacto, neo-Luddite indictment, but did not apologize for his own travels: With the facilities for transportation developed during the past century, migration has reached a stage absolutely unheard of in previous history, and the influx of millions of men into previously unoccupied regions is reflected in immense changes in the vegetation of nearly all parts of the world–far greater than in any previous period of the world's history. Forests have been swept away until the world is menaced with a timber famine, and their place has been taken by crops of all kinds, which are entirely alien to the country and completely alter the appearance of the landscape. This disturbance of the natural vegetation is not confined to the white settler alone, but the natives of many regions, as in Central Africa, and our own western plains materially reduced the extent of forest 89 Charles Elton cited one of Campbell’s later papers in The Ecology of Invasions, but not for its conservation philosophy. 90 Douglas H. Campbell, An Outline of Plant Geography. (New York: Macmillan, 1926) 17. 91 Ibid.,18 iv 48 lands, by cutting and burning–partly to increase the extent of grass-lands for grazing of cattle or wild game. 92 Campbell repeated and presumably believed the pre-colonial “empty America” myth. “The extraordinary and rapid change in the vegetation of a large area, due to man's activities is especially apparent in the United States, which a century ago was to a great extent untouched by man.”93 His xenophytophobia was such that it produced a rare instance in which economically valuable introduced plants were counted as little better than weeds: “Except for the native trees, the predominant vegetation at the present time is largely exotic. None of the staple food crops are indigenous, and the same is true for most of the common fruits, although some of the latter–like grapes and berries of various kinds–are of native origin. Even the weeds are mostly foreigners and have driven out the native woodland plants which have retreated before these hardy invaders.”94 Campbell went on to complain about “escaped” ornamental plants, an idiom attributed to Hooker but uncommonly used until much later.95 Besides the economic plants introduced by man he has imported from all over the world a host of ornamental species which adorn his gardens. Sometimes these escape from cultivation and become quite naturalized. In the cool moist coastal regions of northern California and Oregon one meets the showy broom and foxglove of Europe– and one sees these same plants in the similar climates of central Chile and New Zealand. In the latter country the sweet briar and blackberry, introduced from England 92 Ibid., 18-19 93 Ibid., 19 94 Ibid., 19 95 Oxford English Dictionary Online. iv 49 as garden plants, have escaped and become extremely troublesome weeds.” Most plants ranking as weeds, however, have been introduced accidentally, and wherever man has migrated, weeds have followed him, or been imported from divers sources. Many of the worst weeds owe their rapid dissemination to specially favorable adaptations for seed dispersal, like the wind-borne seeds of thistles and dandelions, or the hooks of burdocks and cockle bur, which stick to the coats of animals or men. Weed seeds may come in mingled with seed grain, or in the dirt adhering to animals or in the cargo of ships and railways. Rapid transit for mankind furnishes equally quick transport for those unwelcome immigrants, and one finds the common weeds of most localities very far away from their original homes. 96 Showing little curiosity about how such things could be, he simply stated, “Many of these introduced plants seem to grow with increased vigor in their new home, and may largely replace the native plants.97 Not content with plants, Campbell added animals to his indictment, but then had to backpedal slightly on crops: “Man has also been responsible for the introduction of many animal pests–injurious insects, rats and mice, rabbits, etc., which may do great damage to vegetation, and greatly affect the flora of large areas. Destructive fungi have also been introduced by human agencies, and are responsible for extensive damage to crops, or native species.”98 Henry N. Ridley Henry N. Ridley was Director of the Botanical Gardens at Singapore, largely responsible for promoting rubber plantations in southeast Asia and author 96 Campbell, Outline (cit. n. 90) 20-21 97 Ibid., 21 98 Ibid., 21 iv 99 of a five volume Flora of the Malay Peninsula. 50 One of the great idiosyncratic descriptions of neobiota appeared within Ridley’s even greater idiosyncratic account of modes and methods titled The Dispersal of Plants Throughout the World. As arguments go, The Dispersal of Plants was discursive and difficult, but also densely packed with information. Its thirty-page chapter “Dispersal by Human Agency” followed suit. Ridley’s citations for concepts were sometimes precise and sometimes vague, making it difficult to tell whether ideas were his own or received. His bibliography was extensive, but did not mention Campbell, whose work may simply have come out after Ridley’s project was essentially complete. Ridley related most of his concepts via examples instead of synthesis. But many seem familiar today. For example: “In Singapore there was, for many years, a patch of an American grass … growing along the roadside near the Botanic Gardens. This was the only known patch of this grass in the Malay Peninsula. It occurred to me to utilize it as a broad-leaved grass for flower-border edges, and I had some dug up, and propagated and planted in the Botanic Gardens. In a few years it had spread, not only all over the Gardens and along the roadsides all over Singapore, but into the Malay Peninsula as far as Selangor. [roughly 200 miles] The original patch had been hemmed in by strongergrowing creeping grasses, so that it had been quite unable to spread; but as soon as it reached a spot where there was little or no competition, it was able to spread, and actually drove out the other grasses.100 99 Edward J. Salisbury, “Ridley, Henry Nicholas (1855–1956).” Rev. Oxford Dictionary of National Biography. Ed. H. C. G. Matthew and Brian Harrison. (Oxford: University Press, 2004). 100 Henry N. Ridley, The Dispersal of Plants Throughout the World. (London: L. Reeve, 1930) 629. iv 51 Ridley did not dwell on generalities; unlike his predecessors, he limited his categories to indigenous and (by default) introduced. “I reserve the word indigenous” he wrote, “for plants which arrived at their present stations by natural methods, by wind, or wild animal transport, or were evolved from ancestors which were brought there by such agencies.”101 One of Ridley’s major phytogeographical concerns was to demolish an existing category of botanical convenience. “Formerly a large number of plants were recorded in botanical works as Cosmopolitan, a word accepted as meaning that they were to be found in any part of the world where the climate and environment were suitable to their growth. I have paid special attention to this class of plants (i.e., those of especially wide distribution), and find that, in flowering plants, the greater number owe their primary distribution to accidental or intentional human transport, though, when they arrived at their new home, they were usually still further dispersed by natural causes.”102 His specific conception of intermediate but essentially prehistoric dispersal by humans might have been an original one. Ridley found that by 1930, “A large number of works and papers [had] been published on the alien plants of different parts of the world, and it would be impossible to give a full account of the innumerable species which have found their way from one corner of the globe to another by the aid of man, and to have 101 Ibid., 628. 102 Ibid., 628. iv 103 established themselves.” 52 Nevertheless, for another thirty pages, he tried to do so. Taking his caveat to heart, I quote him on only one more topic. Again it is hard to know whether he came by this conception via reading or reflection, but it is a valuable one. “A plant whose area of occupation is very limited, by an alteration of the environment of that, or of a near-lying area, whether by human action or natural action, may become extremely abundant and spread over a very large area, and, in fact, from being very rare may become very common. Here man has, in this way, often effected a vast change in the flora, exterminating some species and causing rare plants to become common, and common plants rare.”104 Evgeniĭ Vladimirovich Wulff Crimean-born and Vienna educated botanist E.V. Wulff was curator of the Herbarium of Cultivated Plants in the Geography Department of the Institute of Plant Industry when he was killed during the 1941 Nazi assault on Leningrad.105 Wulff “stated on many occasions that in order to know botany well it is necessary to learn its history,” and demonstrated his commitment to this idea by translating many of Alexander von Humboldt’s works into Russian.106 Wulff’s An Introduction to Historical Plant Geography included a fifteen-page chapter on the “history of the science;” there and elsewhere in his text he demonstrated his 103 Ibid., 630. 104 Ibid., 634. 105 Vladimir Asmous, “Dr. E.V. Wulff—A Victim of War.” Chronica Botanica 9 (1945) 210-211. 106 Ibid. iv 107 familiarity with most of the botanists previously mentioned here. 53 Wulff collected his thoughts on neobiota in a chapter on “artificial factors.” Like many who came before, he distinguished between intentional and accidental introductions, but he also made a point of showing how weeds and crops underwent parallel forms of artificial selection.108 He also echoed the earlier distinction between station and habitation: The significance of man in changing the vegetation of the earth, particularly as regards disturbing its natural state, is unquestionably immense; his role in the distribution of plants over the globe is, no doubt, very considerable, but it is, nevertheless, greatly limited by the specific habitat conditions required by each particular species of plant. The significance of man’s role with respect to the present distribution of species throughout the world has been greatly exaggerated, and it is only during the past few centuries tat man’s role has acquired the importance that it has. Only the comparatively recent, extensive development of ways of communication and the constant movement of people and freight from region to region, as well as the expansion of the cultivated areas and the introduction into cultivation of plants from distant countries, have enabled man to assume such an important role in the geographic distribution of plants.109 Wulff was also unique by virtue of his enthusiasm for Wegenerian continental drift long before its mechanism was accepted: Its significance for biogeography is beyond doubt. It has thrown light on a number of hitherto entirely incomprehensible moments in the geography of plants. It has shown us new ways out from that blind alley into which we were led by the mutually exclusive theories of land-bridges and of the permanence of oceans and continents … even though Wegener’s theory still requires additions and corrections, it, nevertheless, constitutes the 107 E.V. Wulff, An Introduction to Historical Plant Geography, transl. Elizabeth Brissenden. (Waltham, MA: Chronica Botanica, 1943). 108 Ibid.,104-106. 109 Ibid., 113-114. iv 54 only plausible working hypothesis upon which the historical plant geographer may base his conclusions.110 The Zoologists It took zoologists quite a while longer than the botanists to develop an organized interest in neobiota, and once they did, many included botanical examples and elements in their discussions. Even so, there remained one notable and obvious difference between the botanical and zoological literatures. The zoologists never really warmed to the task of anekeitaxonomy. Any of the categories discussed above could have been adapted to animals, but this apparently didn’t happen. Perhaps the relatively smaller (known) number of neobiotic animal species obviated the need for categories, or the many observational difficulties of zoological study rendered the problem insoluble. So far, I have encountered no examples of such attempts, no references to any, and no mentions of any such need during the period under consideration. This section covers several authors (but only one really familiar name) whose work was published between 1851 and 1950. Some were specialists, others hardly so, but all generalized about neobiota to some extent. They are organized by their major interests, which may be taxonomic (e.g., mollusks) or geographic (e.g., New Zealand) and within those categories they appear in order of original publication. Again, I am dealing only with major accounts. 110 Ibid., 199. iv 55 Binney and Kew: The Conchologists Nineteenth century conchology (later, malacology) was rife with discovery and discussion of neobiotic bivalves and gastropods. Being relatively sessile, at least as adults, neobiotic shells might easily have inspired anekeitaxonomies, though none have been identified. However, two comprehensive accounts (one from the United States and the other British) stand out among the incidental chatter of published journal papers and letters. Born in 1803, Bostonian Amos Binney graduated from Brown University at seventeen, and began studying medicine. His own health was precarious, but he managed to complete a Harvard M.D. by 1826. He was evidently fascinated by land snails, and devoted his time and eventually his self-made fortune to collecting and cataloguing them. Binney co-founded the Boston Society of Natural History, serving as Curator, founding its first journal, and finally, as President of the organization. He died at forty-four, leaving an unfinished magnum opus.111 This material was published posthumously (between 1851 and 1857) as The Terrestrial Air Breathing Mollusks of the United States and the Adjacent Territories of North America. Like some others discussed in this chapter, the title of Binney’s series belied its contents. Volume one (of five) opened with three chapters titled: “Of the rise of scientific zoölogy in the United States, with notices of naturalists particularly connected with the subject of this work,” “Of the ignorance and 111 Richard I. Johnson, “The Rise and Fall of the Boston Society of Natural History” Northeastern Naturalist (2004): 81–108. iv 56 neglect of American labors in zoölogy exhibited by European naturalists,” and then “Of some of the obstacles impeding the study of zoölogy, and the means of overthrowing them.”112 But chapters five through eight dealt with distributions, including many of what we now label ecologically ‘limiting factors.’ Binney died a dozen years before Darwin published The Origin and there is no telling what he would have made of natural selection. His avowed conception of creation dutifully followed the biblical line, but with very thinly veiled reservations. After giving an account of the strange distribution of Helix pulchella, and noting in particular its 1820 collection in mid-continent North America, he speculated: Its existence in this case involves some important consequences, for, if we consider it to be of foreign origin, and take the period of its introduction to have been some time since the first permanent colonization of this continent by Europeans, and suppose the point at which it was introduced to have been upon the sea-coast, it is necessary that the animal should have travelled more than twenty-thousand times its own length every day while in motion, and to have been in progressive motion one-fourth of the whole time for two hundred years, in order to have reached this locality; and if its progress had been aided by accidental transportation to some point on the Mississippi River, the result will not be the less improbable … It is true, that the historical period within which this continent has been known, comprises probably but a small part of its whole existence, and that the action of known causes through a period of indefinite duration, might have produced even a more general diffusion, but, as an impenetrable veil hangs over everything that preceded the historical epoch, and we know of no facts which corroborate this latter suggestion, we cannot place much reliance upon it. We must then seek for other causes, to explain the general dispersion of this and other cosmopolite species. Of the origin and mode of creation of organized beings, we of course can know nothing, through our own limited faculties. The subject is beyond our comprehension, and Divine Providence has vouchsafed to us no revelation concerning it. The Mosaic account of creation informs us that 112 Evidently, even an invertebrate specialist may have bones to pick. iv 57 after the surface of the earth was prepared for the support of animal life, the different classes of animals were created at different periods of time, and our own experience, drawn from observation of the fossil remains of former animals, which have been preserved in the strata of the earth’s crust, fully corroborate this account. But, we are limited to these very general facts, and must found our views of the local origin, and the subsequent dispersion of species over the earth upon such observations as we possess, and such reasonings as we can base upon them.113 Binney, in the now-familiar pattern, went on briefly to discuss centers of creation. He faulted zoologists for devoting their interests in this regard to “large animals, and almost solely to the vertebrata.”114 His conclusion: “we find no difficulty in supposing that under the same or similar conditions, the same species may have been created at different centres. In this way the presence of any species in every part of the earth may be accounted for, and thus only can we satisfactorily explain the diffusion of the species that have been under consideration.”115 This seems to preclude much further meaningful discussion of specific origins and modes of distribution, but apparently not so to Binney’s mind. Half a century after Amos Binney’s death, English “amateur naturalist” Harry Wallis Kew published a molluscan foreshadowing of Ridley’s Dispersal of Plants as a volume in the respected and successful “International Scientific Series.” Little biographical information has come to light regarding Kew. One 113 Amos Binney, The Terrestrial Air Breathing Mollusks of the United States and the Adjacent Territories of North America (1851) I, 143-144. 114 Ibid., 145. 115 Ibid., 148-149. iv 116 source reported that he was a bank clerk in Kent. 58 His book’s footnotes suggest that he was a regular reader of numerous journals and magazines devoted to science and natural history. Like Binney’s earlier effort, it concentrated on terrestrial species, but added freshwater aquatics as well. Simply titled The Dispersal of Shells, Kew’s book carried an auspicious preface by Alfred Russell Wallace, contextualizing the work’s contribution to the general inquiries of zoogeography. Alerting readers to Kew’s method and manner, Wallace wrote: Many books of far greater pretension, even though they should contain descriptions of scores of new species and work out their internal structure with the greatest accuracy, may yet be of less interest to the philosophical naturalist than this unpretending little volume. In its pages we are afforded a glimpse of what seem at first sight to be but trifles and accidents in nature’s workshop, but which are really the tools with which she produces some of her most striking results. It is owing to such trifling occurrences as the attachment of a living shell to a beetle’s leg, or the conveyance of seeds in the mud on a bird’s foot, that many remote islands have become stocked with life, and the range of species extended or modified over the earth; while through changes of the organic environment thus effected even the origination or the extinction of species may have been brought about.”117 Editor John Carrington reviewed The Dispersal of Shells for his naturalists’ magazine Science-Gossip, and his comments are more usefully revealing than any quotations from the book itself. “Without encumbering his pages with useless speculation, the author has gathered together a most useful mass of facts that form one of those interesting stories which we too seldom find 116 Hans G. Hansson, Biographical Etymology of Marine Organism Names (BEMON) Tjärnö Marine Biological Laboratory (1997) http://www.tmbl.gu.se/ accessed 13 October 2006. 117 Alfred R. Wallace, “Preface,” to Harry Wallis Kew, The Dispersal of Shells (London: Kegan Paul, 1893), v-viii. iv 59 associated with books of scientific value.” Carrington also drew attention to Kew’s 86-page closing description of neobiota: “Chapters viii. and ix. are well worth attention, as showing the influence of the civilisation of mankind in unintentionally contributing to the equalization of the fauna of various regions, if climatically suitable. This agency must necessarily increase in force now that travel and transit has become so rapid and certain with the aid of steam.”118 But Carrington noted, too that Kew had abstained from more than just useless speculation. “The most disappointing defect in this work is the want of a final summary attached to each chapter, of the evidence and conclusions to be drawn, which would have added greatly to the value of the work. We can commend the author’s modesty in speculation, but these conclusions from one who has had so great experience in investigating the subject, could not have failed to interest.”119 Perhaps Kew drew his own line between natural history and biology “not wisely, but too well” for Carrington’s taste. The Motile Taxa: General Zoogeography In 1858 ornithologist Philip Lutley Sclater opened a paper for the Linnaean Society with a surprising premise: “An important problem in Natural History, and one that has hitherto been too little agitated, is that of ascertaining the most natural primary divisions of the earth's surface, taking the amount of similarity or 118 John T. Carrington, “Books to Read” Science-Gossip [n.s.] 1(1893), 15-16. 119 Ibid. iv 120 dissimilarity of organized life solely as our guide.” 60 It was surprising because rather a lot of work by botanists had gone into this question during the preceding half century. But ignorance of earlier work, especially outside an author’s specialized interests, is a hallmark of this topic. It is easy to imagine as an aspect of later science, and the proliferation of narrowly-focused journals; it is more difficult to imagine in the days when “natural history” was still considered a major disciplinary focus. Sclater made his position and purpose plain: “It is a well-known and universally acknowledged fact that we can choose two portions of the globe of which the respective Faunæ and Floræ shall be so different, that we should not be far wrong in supposing them to have been the result of distinct creations. Assuming then that there are, or may be, more areas of creation than one, the question naturally arises, how many of them are there, and what are their respective extents and boundaries, or in other words, what are the most natural primary ontological divisions of the earth's surface?”121 What then emerged was a division of the planet into six ornithological realms, forming the basis of much later zoogeographical work, and surviving (under another name) to strongly influence Charles Elton. Sclater dismissed the idea that zoogeographical dynamics occurred without human influence: “We do not find that the Nightingale extends its range farther to the west one year than 120 Philip Lutley Sclater, “On the General Geographical Distribution of the Members of the Class Aves,” Journal of the Proceedings of the Linnean Society: Zoology 2 (1858), 130-145 121 Ibid. iv 61 another, nor that birds looked upon as occasional visitors to this country, grow more or less frequent. If the contrary be the case, it may always be accounted for by some external cause, generally referable to the agency of man, and not to any change in Nature's unvarying laws of distribution.” He also found multiple centers of creation appealing for another purpose: I suppose few philosophical zoologists, who have paid attention to the general laws of the distribution of organic life, would now-a-days deny that, as a general rule, every species of animal must have been created within and over the geographic area which it now occupies. Such being the case, if it can be shown that the areas occupied by the primary varieties of mankind correspond with the primary zoological provinces of the globe, it would be an inevitable deduction, that these varieties of Man had their origin in the different parts of the world where they are now found, and the awkward necessity of supposing the introduction of the red man into America by Behring's Straits, and of colonizing Polynesia by stray pairs of Malays floating over the water like cocoa-nuts, and all similar hypotheses, would be avoided.”122 Forty years after outlining his six-way terrestrial division, Sclater proposed six “sea-regions” for the geography of marine mammals. He acknowledged but could not account for the failure of south Atlantic species to spread northwards, suggesting that “in former ages there must have been some barrier in the Atlantic … the only barrier I can imagine that would have effected this must have been a land uniting South America and Africa, across which they could not travel.” Nevertheless, he was not prepared to abandon his basic premise, concluding: “All these facts, with the one exception of the supposed Atlantic Barrier, would tend in favor of the now generally accepted doctrine that the principal masses of 122 Ibid. iv 62 land and water are not of modern origin, but have existed mainly in their present shapes throughout all ages.”123 In contrast with his terrestrial regions, by 1958 Sclater’s sea regions were barely acknowledged except as a failed paradigm of marine mammalogy.124 There is no reason to suppose that all of Sclater’s motivations survived along with his realms in the zoogeographical work of the following century, but it is interesting to ponder how his divisions might have differed given more dynamic assumptions. Sidney I. Smith Sidney Smith was a Yale zoologist, a comparative anatomist of insects, and later of marine crustaceans.125 In 1867, apparently unaware of either Binney or Sclater, he wrote “The Geographical Distribution of Animals.” Another example of narrow specialization or just spotty research; both omissions are puzzling. Smith did cite a paper from the Proceedings of the Boston Society of Natural History (which Binney had founded); Smith did not study shells but he did study invertebrates, and could have been expected to know of major recent works in that regard. In that citation and another, Smith used examples of bird 123 Philip Lutley Sclater, “On the Distribution of Marine Mammals,” Science 5 (1897): 741-748. 124 J. L. Davies, The Pinnipedia: An Essay in Zoogeography Geographical Review 48 (1958) 474-493 125 Anonymous, “Sidney Irving Smith.” Yale Peabody Museum: History and Archives, www.peabody.yale.edu/archives/ypmbios/smith_si.html, accessed 15 October 2006. iv 63 distributions, suggesting that the topic and Sclater’s Ibis had at least come to his attention. As was typical of other biogeographies, Smith’s devoted only a fraction of his discussion to neobiota. And like many others, his had distinctly theological overtones: “Man, with boundless aspirations and governed in all things by an influence within himself, is given a power in nature second only to his Creator; with control over physical causes, he is governed by no laws of geographical distribution, and , traversing the whole earth at his will, he has carried, in spite of climatic influences, species from continent to continent, and almost pole to pole. His influence — far above all other secondary causes, and uncontrolled by the laws imposed upon mere animals — seems only a disturbing force among the naturally harmonizing laws of the diffusion of life.” 126 Smith did display some familiarity with the recent literature of one of his specialties, but he closed this paragraph with an ambiguous comment: Three common species of butterflies in Eastern North America … long known to be identical with European species, have been asserted to be natives of this country, and the possibility of their introduction from Europe has recently been questioned. But within a very few years, there has been a well-authenticated instance of the naturalization of an European butterfly in Canada. [This species] was introduced at Quebec about 1859, and, in 1863, it had become very abundant within a circle of forty miles radius about that city. If butterflies are introduced and spread so rapidly now, there is no reason why the other butterflies [which] feed upon introduced plants, should not have been introduced and diffused over all the eastern part of the country long before entomologists began to study the distribution of species.127 As published in the American Naturalist, Smith’s article bore an editorial note stating “‘The Influence of Secondary Causes in the Geographical 126 Sidney I. Smith, "The Geographical Distribution of Animals." American Naturalist 2 (1868) 124-31. 127 Ibid. iv 64 Distribution of Animals,’ [was] one of the subjects assigned for essays for the Berzelius prizes in the Sheffield Scientific School of Yale College in 1867.”128 So far, no competing entries have come to light, nor have details of the motivation for assigning this subject. The note pointedly does not identify Smith as the prizewinner. Unless the fix was in, the existence of this competition suggests that the topic was interesting to someone at Yale besides Smith himself. Alfred Russell Wallace In 1859, perhaps benefiting from his recent “natural selection” notoriety, Alfred Russell Wallace suggested some revisions to Sclater’s regional boundaries, which Sclater published in Ibis, his own new journal. This began (however unwittingly) the process by which the regions would become known as “Wallace’s Realms.”129 But like Sclater, Wallace barely broached the topic of neobiota in his own massive, 1876 The Geographical Distribution of Animals, still acknowledged as a seminal biogeographical text.130 Neither “man” nor “humans” rated an entry in his index. Species introductions are mentioned tersely, if at all, and some by-then notorious ones were ignored. For example, In the mid-1860s, rabbit populations in Australia were already burgeoning.131 By 1874, Wallace reported, “An Indian [rabbit] species is now wild in some parts of Java, but has probably been introduced,” but failed even to acknowledge the Australian 128 Ibid., 124n. 129 Alfred Russell Wallace, “A Letter from Mr. Wallace Concerning the Geographical Distribution of Birds.” Ibis 1 (1859), 449-454. 130 John C. Briggs and Christopher J. Humphries, “Early Classics,” in Lomolino, et al, Eds, Foundations of Biogeography (see cit. n. 26), 11. 131 David G. Stead, The Rabbit in Australia (Sydney: Wynn 1935), 16-17. iv 132 situation. 65 Similarly, imported house sparrows were already a hot topic of debate in the United States, but Wallace remained silent on this matter, as well. The purported effects of recently distributed domestic animals briefly got his attention: “We know, for example, that the introduction of goats onto St. Helena utterly destroyed a whole flora of forest trees; and with them all the insects, mollusca, and perhaps birds directly or indirectly dependent on them. Swine, which ran wild in Mauritius, exterminated the Dodo.”133 But he did not really differentiate between the effects of domestic animals on wild animals and the effects of wild animals on domestic ones. Elsewhere in the “dodo” paragraph he noted “So, in South Africa, the celebrated Tsetse fly inhabits certain districts having well-defined limits; and where it abounds no horses, dogs or cattle can live.”134 Within five years Wallace took note of neobiota, or at least began to consider them worth mentioning. In Island Life he wrote, for example, that “rabbits, weasels, rats and mice, and a small lizard peculiar to Madeira and Tenerife are now found wild in the Azores, but there is good reason to believe these have all been introduced by human agency,” and went on to include the island’s freshwater fishes among the obviously transplanted species.135 At greater length he discussed the advent of European and Australian mammals in 132 Wallace, The Geographical Distribution of Animals (London: MacMillan,1876) II, 242-243. 133 Ibid., I, 44. 134 Ibid., I, 45. 135 Alfred Russell Wallace, Island Life, (New York: Harper, 1881), 236. iv 66 New Zealand, and eloquently decried the exploitation and eventual replacement of the indigenous flora and fauna of St. Helena.136 But none of these examples led him to generalize about neobiota in Island Life’s “Summary and Conclusions,” or even to predict that similar events would unfold on other islands. Later still, in Darwinism, Wallace finally ventured to generalize. Since his remarks constituted an early assertion of relationships among different cases of neobiota, and explicitly relied on now-familiar predecessors, it seems appropriate to represent him in his own words. [A] phenomenon of an analogous kind [to weeds gradually excluding garden plants] is presented by the different behaviour of introduced wild plants or animals into countries apparently quite as well suited to them as those which they naturally inhabit. Agassiz, in his work on Lake Superior, states that the roadside weeds of the northeastern United States, to the number of 130 species, are all European, the native weeds having disappeared westwards; and in New Zealand there are no less than 250 species of naturalised European plants, more than 100 species of which have spread widely over the country, often displacing the native vegetation. On the other hand, of the many hundreds of hardy plants which produce seed freely in our gardens, very few ever run wild, and hardly any have become common. Even attempts to naturalise suitable plants usually fail; for [Alphonse] de Candolle states that several botanists of Paris, Geneva, and especially of Montpellier, have sown the seeds of many hundreds of species of hardy exotic plants in what appeared to be the most favourable situations, but that, in hardly a single case, has any one of them become naturalised. Even a plant like the potato—so widely cultivated, so hardy, and so well adapted to spread by means of its manyeyed tubers—has not established itself in a wild state in any part of Europe. It would be thought that Australian plants would easily run wild in New Zealand. But Sir Joseph Hooker informs us that the late Mr. Bidwell habitually scattered Australian seeds during his extensive travels in New Zealand, yet only two or three Australian plants appear to have established themselves in that country, and these only in cultivated or newly moved soil.137 136 Ibid., 278-81, 437. 137 Alfred Russell Wallace, Darwinism, (New York: Macmillan, 1889), 15-16. iv 67 He revisited St. Helena, and then the Middle East: The introduction of goats into the island of St. Helena led to the entire destruction of the native forests, consisting of about a hundred distinct species of trees and shrubs, the young plants being devoured by the goats as fast as they grew up. The camel is a still greater enemy to woody vegetation than the goat, and Mr. [George Perkins] Marsh believes that forests would soon cover considerable tracts of the Arabian and African deserts if the goat and the camel were removed from them. Even in many parts of our own country the existence of trees is dependent on the absence of cattle.138 Wallace also used examples from Russia and Australasia to exemplify a phenomenon that would later be called “competitive exclusion”139 : The black rat (Mus rattus) [sic] was the common rat of Europe till, in the beginning of the eighteenth century, the large brown rat (Mus decumanus) [sic] appeared on the Lower Volga, and thence spread more or less rapidly till it overran all Europe, and generally drove out the black rat, which in most parts is now comparatively rare or quite extinct. This invading rat has now been carried by commerce all over the world, and in New Zealand has completely extirpated a native rat, which the Maoris allege they brought with them from their home in the Pacific; and in the same country a native fly is being supplanted by the European house-fly. In Russia the small Asiatic cockroach has driven away a larger native species; and in Australia the imported hive-bee is exterminating the small stingless native bee. The reason why this kind of struggle goes on is apparent if we consider that the allied species fill nearly the same place in the economy of nature. They require nearly the same kind of food, are exposed to the same enemies and the same dangers. Hence, if one has ever so slight an advantage over the other in procuring food or in avoiding danger, in its rapidity of multiplication or its tenacity of life, it will increase more rapidly, and by that very fact will cause the other to decrease and often become altogether extinct. In some cases, no doubt, there is actual war between the two, the stronger killing the weaker; but this is by no means necessary, and there may be cases in which the weaker species, physically, may prevail, by its power of more rapid multiplication, its better withstanding 138 Ibid., 17. 139 See also Garrett Hardin, “The Competitive Exclusion Principle” Science 131 (1960) 1292-1297. iv 68 vicissitudes of climates, or its greater cunning in escaping the attacks of the common enemies (…). As an effect of this principle, we seldom find closely allied species of animals or plants living together, but often in distinct though adjacent districts where the conditions of life are somewhat different … of course there are also numbers of closely related species in the same country, but it will almost always be found that they frequent different stations and have somewhat different habits, and so do not come into direct competition with each other; just as closely allied plants may inhabit the same districts, when one prefers meadows the other woods, one a chalky soil the other sand, one a damp situation the other a dry one. With plants, fixed as they are to the earth, we easily note these peculiarities of station; but with wild animals, which we see only on rare occasions, it requires close and longcontinued observation to detect the peculiarities in their mode of life which may prevent all direct competition between closely allied species dwelling in the same area.140 Still later in the volume, Wallace “realm-hopped” again, among other things, stating one form of a hypothesis that Charles Elton would investigate a half-century afterward: Divergence of character has a double purpose and use. In the first place it enables a species which is being overcome by rivals, or is in process of extinction by enemies, to save itself by adopting new habits or by occupying vacant places in nature. This is the immediate and obvious effect of all the numerous examples of divergence of character which we have pointed out. But there is another and less obvious result, which is, that the greater the diversity in the organisms inhabiting a country or district the greater will be the total amount of life that can be supported there … [t]he same principle is seen in the naturalisation of plants and animals by man's agency in distant lands, for the species that thrive best and establish themselves permanently are not only very varied among themselves but differ greatly from the native inhabitants. Thus, in the Northern United States there are, according to Dr. Asa Gray, 260 naturalised flowering plants which belong to no less than 162 genera; and of these, 100 genera are not natives of the United States. So, in Australia, the rabbit, though totally unlike any native animal, has increased so much that it probably outnumbers in individuals all the native mammals of the country; and in New Zealand the rabbit and the pig have equally multiplied.141 140 Wallace, Darwinism, (cit. n. 137) 34-35. 141 Ibid., 110. iv 69 Wallace used neobiota to make quite a few further points about evolution by natural selection in Darwinism. And he left no doubt that he (like Hooker) felt that examples of wholesale biotic revision like that occurring on St. Helena were deplorable. With Darwinism, Wallace also conspicuously distanced himself from Darwin’s own assertions that humans had evolved like other animals. But he issued neither a blanket endorsement, nor a general indictment, of neobiota. James Ritchie One might expect a book titled The Influence of Man on Animal Life in Scotland: A Study in Faunal Evolution (1920) to be fairly technical and perhaps narrowly scoped. Alas for Ritchie, his book’s title alone may have cost him a place of repute among the earliest proponents of invasion biology. He did not limit his geographical interest to Scotland; rather, he often used Scottish issues as points of departure, comparing and contrasting local cases and situations with those occurring elsewhere. An inventory of the book’s index turned up about thirty-six references to the United States, and at least thirty to countries in continental Europe; eighteen to New Zealand, fourteen each to Australia and Canada, twelve to African and ten to Asian countries, along with individual mentions of less well-visited (at the time) places such as Mauritius, Mexico, the Falkland Islands and the Panama Canal. Oddly, there were no index entries at all for evolution or for natural selection. Darwin rated three mentions, but of those, two dealt with pigeons.142 142 James Ritchie, The Influence of Man on Animal Life in Scotland: A Study in Faunal Evolution (Cambridge, UK: Cambridge University Press 1920), 518-550. iv 70 Perhaps three-quarters of Ritchie’s Influence was devoted to discussing faunal conservation concerns such as deforestation, (over)hunting, urbanization, farming, and appropriate river management. But two chapters, and parts of a third, were explicitly devoted to neobiota. Chapter five, “The Deliberate Introduction of New Animals” occupied sixty pages, divided into sections on introductions for utility, sport and amenity. Chapter seven, section three was devoted to “Effects upon the spread of animals of canals, of roads and bridges, and of railways.” Chapter eight, “Camp-followers of Commerce, or Animals Introduced Unawares” required another fifty; its sections dealt with “hangers-on” (e.g., parasites), “stowaways on ships,” “skulkers in dry [stored] food materials,” “foundlings amongst fruit,” “creatures conveyed by plants and vegetables” including nursery stock, and “timber transportees.”143 Exemplifying his extraCaledonian interests, Ritchie also summarized a November 1916 report of import quarantine inspections for the State of California, listing “close on half a hundred different kind of insects caught attempting to run the blockade of the Californian quarantine … one tiny stream in the world’s great ocean of commerce.”144 Strongly attracted to practical issues, Ritchie devoted a later (1931) book to Birds and Beasts as Farm Pests, in which human agency was mentioned only as it applied to particular cases like rats, muskrats, and grey squirrels. But not long after the publication of Birds and Beasts, Ritchie was invited to speak to the BBC radio audience. In April and May 1932 he delivered a six-part series called 143 Ibid., ix-xii. 144 Ibid., 474-475. iv 71 The Changing Face of Nature, which summarized many elements from both books. In episode four (“The War Upon Wild Animals”), Ritchie called the advent of domestic grazing animals in Britain a “terrific invasion of aliens,” terminology Charles Elton echoed, knowingly or not, almost exactly a year later in The Times.145 Ritchie’s final BBC talk, titled “Gains and Losses in Animal Life,” closed out the series with discussions of the unforeseen consequences attending introductions to Britain of Little Owls from Holland and grey squirrels from North America; mentions of the iconic Australian rabbit disaster, and of English sparrows and starlings in America; and a warning about the effects of American muskrats in Europe, and their still-uncertain future in Britain itself.146 Well before Elton, Ritchie was avowedly wary of species introductions, and an enthusiast of nature conservation. And well before Elton, he credited and blamed natives and aliens alike with respect to their useful, troublesome and attractive qualities. Finally, Ritchie very nearly anticipated in full (and even advanced) the conservation philosophy developed by Aldo Leopold and Charles Elton (see chapter two). In closing the first chapter of Birds and Beasts, Ritchie wrote: The pestilence of an animal often varies with the point of view … conflict of interests crops up frequently, sometimes it is intense, occasionally it is of far-reaching importance. No one can doubt that the farmer would benefit by the reduction in numbers of many abundant seed-eating birds and [rodents] … a different point of view is that the naturalist and particularly the bird-lover. Rightly he wishes to see as many and as great variety of wild creatures as the country can support, but he has no right to 145 James Ritchie, “The War Upon Wild Animals” The Listener 7 (1932), 666-667. 146 James Ritchie, “Gains and Losses in Animal Life” The Listener 7 (1932), 756757 iv 72 insist that the aesthetic pleasures of the country-side should be preserved solely at the expense of the farmer. Where a clash occurs between the preservation of wild creatures, beasts or birds, and the interests of a fundamental industry … the industry must have first consideration, and other means found for protecting the wild things whose existence is threatened. Such may be the reasonable reduction in numbers of the culprits, the calling off of slaughter by offering compensation for damage done, or, most satisfactory of all, the creation of special reserves, where beasts and birds can live free from all interference.147 George M. Thomson New Zealand’s post-colonial national identity is bound up with its sense of geographical isolation and biological uniqueness. The perceived loss of that uniqueness was the sole topic of three more major studies of neobiota in the next century. The first of these, George M. Thomson’s The Naturalisation of Animals & Plants in New Zealand obviously did not limit itself to animals, and calling Thomson a zoologist or his work strictly zoological is thus problematic. But his major focus on neobiota was faunal. He bracketed The Naturalisation with two slimmer volumes of derivative juvenilia devoted to Wild Life in New Zealand. Part one (1921) dealt with mammals; all the mammals in New Zealand excepting pinnipeds and bats were introduced. In many respects, The Naturalisation is part one’s expanded, “grown-up” successor, although within part one the larger volume is already listed as “now in the press.”148 Eventually, Wild Life part two 147 James Ritchie, Birds and Beasts as Farm Pests (Edinburgh: Oliver and Boyd, 1931), 5-6. 148 George M. Thomson, Wild Life in New Zealand, Part I. Mammalia (Wellington: New Zealand Board of Science and Art, 1921), 2. iv 73 (1926) addressed introduced birds and fishes, to “help the young people to a better knowledge of the natural history of their beautiful country.”149 The Naturalisation was undertaken with unshakable confidence and encyclopedic intent: “The naturalisation of animals and plants in any country is a most interesting and fascinating subject, as well as being one of very great and far-reaching importance. In the present work I have endeavoured to state what is known of the subject, as far as it relates to New Zealand. I have stated the facts regarding the first introduction of every species into the country, as far as these can be ascertained, and its subsequent success or failure in establishing itself.” With equal confidence, Thomson continued, “In gathering the information required and working out the material, it was soon evident to me that the subject was unique. It had never been attempted before– as far as I am aware– for any country. Indeed it was seen that New Zealand was the only country in which such a bit of history could be attempted with any prospect of success.”150 Thomson was ignorant at best regarding the uniqueness of his vision, given (e.g.,) H.C. Watson’s work on British plants and Ritchie’s on Scottish animals. But his point about potential success was better taken. Although New Zealand entered European consciousness in 1642 with the arrival of Dutch explorer Abel Tasman, it was essentially ignored for a further century until coastal explorations of the islands began under James Cook in 1769. Whaling 149 George M. Thomson, Wild Life in New Zealand Part 2. Introduced Birds and Fishes (Wellington: New Zealand Board of Science and Art,1926), 5. 150 George M. Thomson, The Naturalisation of Animals & Plants in New Zealand (Cambridge, England: Cambridge University Press 1922), 1. iv 74 and missionary posts appeared by the 1790s, but colonization did not begin in earnest until 1839.151 The relatively late and well-organized arrival of Europeans, their plants, animals and technologies in New Zealand (by comparison with the North America, for example) was also better documented. Many introductions were undertaken by dedicated acclimatisation societies, and most of them maintained records of their activities. Thomson wrote, “the missing records and blank pages are very numerous,” but “we possess a fairly accurate record of what was here when the Europeans first visited these shores, and we have been able to follow the later introductions of new species with a certain measure of success.”152 Thomson was a seemingly hyperkinetic educator and natural historian. As one of the less well-known names in this chapter, his biography merits a brief digression. 153 A child of empire, Thomson was born in Calcutta to Scottish parents and then educated in Edinburgh. The family eventually relocated again, to New Zealand. They failed there as farmers, but George, with some university training in botany and chemistry, began tutoring at a Dunedin high school. He went on to found colleges and kindergartens, and publish “375 scientific articles, mainly on plants, crustaceans, fish and fisheries,” discovering and describing the 151 Angus J. Harrop, England and New Zealand from Tasman to the Taranaki War (London: Methuen 1926). 152 Thomson, The Naturalisation, (cit. n. 150) 1. 153 Except as cited in the following note, all biographical information for Thomson in this paragraph is from, E. Yvonne. Speirs 1993. “Thomson, George Malcolm 1848 - 1933.” Dictionary of New Zealand Biography, updated 7 April 2006; accessed online 16 October 2006 at http://www.dnzb.govt.nz/ iv 154 crustacean family Anaspididae in 1894. 75 He successfully proposed the construction of a marine biological station, and served on the board of the Portobello fish hatchery, where he also researched acclimatization methods. Thomson is credited with instituting significant scientific reforms in New Zealand, and even served briefly in Parliament. He married twice and fathered six children, outliving both wives and all but two of his offspring. In his idle hours, he was a Presbyterian elder, deacon, and occasional organist, and sang with the Dunedin Choral Society. Much of this he accomplished while standing on one foot, having been accidentally shot in the other one at age 34, finally losing the ruined appendage altogether eleven (presumably uncomfortable) years later. The Naturalisation opened with a historic account that set the tone for the entire volume by extensively quoting primary sources. Over eighty percent of the volume’s 600+ pages were occupied by species accounts, arranged in more or less taxonomic order. These opened with charismatic megafauna, “reversing” the animals to begin with mammals, and end with leeches. Plants followed, arranged in a more typical fashion. These accounts covered some 1,200 species, and ranged from several words to several pages long. Anything Thomson found mentioned as having been present in or brought to New Zealand rated an account, even when the organism could not be positively identified according to the available description. Throughout, he did not suffer fools gladly, and made pointed (and often humorous) comments in the face of dubious 154 S.J. Brands, (comp.) “Systema Naturae 2000.” (Amsterdam, 1989-2005) [http://sn2000.taxonomy.nl/] accessed 4 January 2006. iv 76 anecdotes or attempts to evade responsibility for ill-conceived introductions, whether failed or all-too successful. Thomson closed The Naturalisation with several short chapters, two holding particular interest here. Chapter Thirteen he titled “Interaction of Endemic and Introduced Faunas.” Structurally, it consists of an ecological discussion, parenthetically embedded in an evolutionary one. His evolutionary revelation was that he had long expected, but failed ever to observe, evidence of rapid evolutionary change in populations of species introduced to New Zealand. A twenty-first century reader quickly realizes that the evolution Thomson looked (and seemingly longed) for was going to occur rarely at best; watching for readily observable increases in albinism and other color changes in mammals and birds seemed to weigh on him to the exclusion of all other possibilities. For example, he conspicuously did not speculate that the weka, a “chicken-sized” pre-colonial, flightless species of rail, might be evolving in terms of food and habitat preferences and hunting behavior, even though he repeatedly mentioned it as a significant predator on introduced birds and rodents. But Thomson was a busy man, not likely to spend his time cataloguing fine measurements on numerous specimens, and his monopodal configuration precluded much real fieldwork. Thomson’s ecological speculations led him further than his evolutionary ones, and bear striking resemblances to hypotheses under modern discussion. Rhetorically raising the question “What effects have been produced on the native fauna by the introduction of foreign animals and plants onto the islands of the New Zealand group?” he answered himself cautiously, “ the effects have been so iv 77 far-reaching and so complex that it is impossible to present any summary of them, and all that can be done is to show various aspects of the problem, and to consider facts in detail.” But once having offered the requisite excuses for the fragmentary information at his disposal, he was unafraid to make his cases. Thomson listed many examples of dual and multi-species interactions between introduced and pre-colonial species that led to expansion of one population (not always the newcomers) at the apparent expense of others (not always the old guard). But he was convinced that many supposed cases of direct impacts on pre-colonial species by introduced ones were misperceived, and that multiple causes and effects were at play: It must not be supposed that it is the introduced animals alone which have produced [the retreat of the natives], even though rats, cats, rabbits, stoats and weasels, as well perhaps as some kinds of introduced birds, have penetrated beyond the settled districts. It is largely the direct disturbance of their haunts and breeding places, and the interference with their food supply, which has caused this destruction and diminution of the native fauna … [m]any insects which were common in the bush fifty years ago must have been displaced and largely disappeared. I cannot appeal to figures, but the surface burning of open land which prevailed, especially in the South Island, and the wanton destruction and burning of forest which has marked so much of the North Island clearing, must have destroyed an astonishing amount of native insect life, and made room for introduced forms. The clearing of the surface for cultivation and grazing, the draining of swamps, and the sowing down of wide areas in European pasture plants, have all contributed to this wholesale destruction and displacement of indigenous species. The disappearance of mosquitoes and sand-flies in settled district where they were formerly very common is a case in point;– where they disappeared, it is tolerably certain other species not so well known became scarce at the same time.155 Chapter fourteen of The Naturalisation was titled “Alteration in Flora Since European Occupation of New Zealand.” It began with a rapid-fire listing of 155 Thomson, The Naturalisation, (cit. n. 150) 507-8. iv 78 interactions between pre-colonial plants and post-colonial animals, including vertebrate and invertebrate herbivory, granivory, frugivory and seed dispersal, and trampling. Following that was a very different subsection discussing the “Inter-relation of Native and Introduced Flora.” There, Thomson employed several extended quotations from Wallace, Darwin, and Joseph Hooker, as well as his own contemporary countrymen Leonard Cockayne, T.F. Cheeseman and others, in an effort to refute the belief that introduced vegetation would inevitably drive pre-colonization species to extinction.156 The section contains little text that can be considered purely Thomsonian, but one remark stands out: The opinion of all botanists in New Zealand to-day [sic] is that when the direct, or — to a large extent — the indirect influence of man is eliminated, the native vegetation can always hold its own against the introduced. Those plants which have thriven abnormally in this new country, and have impressed visitors by their abundance, are found in settled and cultivated districts, and belong chiefly to what are known as weeds of cultivation, that is, plants which have become adapted to conditions caused by the direct and indirect actions of human beings, and which only thrive where those conditions are maintained.157 Eighty years on, it is easy to question what Thomson could have meant by “direct” and “indirect” actions, but as mentioned above, his overall tone suggests considerable confidence that all the important factors of these cases were readily observable and understandable. To Thomson, “direct action” probably meant physical attacks made by humans on plants (uprooting, girdling, etc.), leaving all other conceivable effects in the “indirect” category. 156 Thomson acknowledged the help of Cockayne and Cheeseman in his preface (Ibid., vii-viii). 157 Ibid., 527-528. iv 79 Kazimierz Antoni z Granowa Wodzicki The most recent of the zoologists to be discussed here was K.A.G. Wodzicki, scion of Polish nobility. Misfortunes recalling Thomson’s, (in this case, the Soviet occupation of Poland and his father’s exile to Siberia) prompted Wodzicki’s resettlement in New Zealand.158 By contrast with Thomson, he arrived with a doctoral degree , and came from a family with a tradition of natural history studies. And by contrast with James Ritchie, Wodzicki focused his book, Introduced Mammals of New Zealand (1950), almost as narrowly as could be expected from the title.159 His introduction provided summary information regarding introduced birds, but the body of the work is wholly mammalian. Wodzicki was the only author identified during this project to have cited Charles Elton’s 1943 “The Changing Realms of Animal Life” before Elton himself wryly drew attention to the paper in his preface to The Ecology of Invasions. Wodzicki even incorporated Elton’s title into his own text, suggesting the little polemic’s influence on his thinking: “it seems comparatively easy to mark the onset of this period in the changing realm of animal life in New Zealand.”160 158 Peter Bull, “Wodzicki, Kazimierz Antoni z Granowa 1900 - 1987”. Dictionary of New Zealand Biography updated 7 April 2006; accessed online 16 October 2006 at http://www.dnzb.govt.nz/. 159 Wodzicki’s written English, and/or that of his editorial assistants, was certainly on par with Elton’s. But the title Introduced Mammals OF New Zealand is slightly paradoxical; so labeled, these taxa seem to acquire at least a grudgingly acknowledged belonging. 160 Wodzicki, Introduced Mammals of New Zealand (Wellington: New Zealand Department of Scientific and Industrial Research,1950) 4. iv 80 An expatriate Polish zoologist could be expected to notice a paper published in a journal aimed squarely at expatriate Polish scientists. But Wodzicki demonstrated comprehensive awareness of previous work. He drew on several Elton publications, including Animal Ecology and Voles, Mice and Lemmings; on both books by James Ritchie, several works by George Thomson, and Aldo Leopold’s Game Management. Introduced Mammals of New Zealand reviewed well. Charles Elton complimented both the author, “with a record of distinguished ecological work in Poland,” and the volume: “Although this book does not pretend to enter into a deep analysis of the ecology of any of these species of mammal,” Elton wrote, “it gives an extremely clear and interesting account of their present status, and is informed by an ecological outlook.”161 In The Ecology of Invasions Elton also reproduced three of the nicely drawn figures that Wodzicki attributed to “Miss Nancy A. Cooper and Mr. Peter C. Bull.”162 In a review for Ecology titled “Tampering with Nature in New Zealand” Wilderness Society (USA) President Olaus Murie enthusiastically summarized Wodzicki’s major points. During his prior tenure at the U.S. Biological Survey, Murie had been an early and outspoken critic of his agency’s “hate-ridden” native predator control programs (and a sympathetic correspondent of Aldo 161 Charles Elton, “Mammals in New Zealand,” The Journal of Animal Ecology, 21 (1952) 161-162. 162 See Wodzicki, Mammals, (cit. n. 160) figs. 1, 41,42; Charles S. Elton, The Ecology of Invasions by Animals and Plants (London: Methuen, 1958) 88, 90, 91. Peter Bull is cited above as Wodzicki’s biographer. iv 163 Leopold’s). 81 Here, he took no issue with controlling introduced species by the familiar methods Wodzicki described. But Murie did pause briefly to snipe at a target closer to home: “Perhaps it is appropriate to mention that the United States government has even at this late date launched an ambitious program of introducing exotic game species into this country, and has personnel touring the world to seek species attractive to American sportsmen.”164 Wodzicki’s account seems relatively objective, especially insofar as he concentrated on analyzing numeric or pseudonumeric data — and he usually called attention to the tenuousness of the latter. Nevertheless, cultural and perhaps idiosyncratic value judgments abounded. He favored species that could be shown to have benefits to society, and since his book is also a government report, may have felt obligated or been persuaded to do so. For example, hares fared better in his view than rabbits. In a few instances, Wodzicki equivocated regarding the status of a beast. He considered wild pigs to be locally helpful in controlling European rabbits and bracken fern, even though they were known to prey on sheep elsewhere.165 Wodzicki felt that cats occupied “a rather peculiar position in New Zealand wild life,” some “domestic”, some “wild,” and others “having severed their connections with Man but have not yet acquired the characteristics of feral animals.” Stating “the degree of domestication has, 163 Curt Meine, Aldo Leopold: His Life and Work (Madison: University of Wisconsin Press 1988), 286-7 164 Olaus J. Murie, “Tampering with Nature in New Zealand,” Ecology 32 (1951) 767-768. As evidenced in his book, Wodzicki’s attitude toward sport hunters and hunting was also fairly deprecatory. 165 Wodzicki, Mammals, (cit. n. 160) 233-4. iv 82 however, only a relative effect on the feeding habits of the cat” provided little elucidation of his point, especially when he followed it with “unless otherwise stated, no discrimination will be made … between wild cats and those with varying degrees of domestication.”166 Animal Anekeitaxonomy? Having previously declared the nonexistence of formal zoological anekeitaxonomies, it remains possible to show evidence for the existence of informal ones. Wallace (1876) and Thomson arranged their discussions phylogenetically, so causal attributions regarding distributions were approached taxon by taxon. In 1881, Wallace called Part I of Island Life “The Dispersal of Organisms, Its Phenomena, Laws and Causes” without really addressing neobiota; Part II he organized geographically. In Darwinism Wallace employed various terms like “naturalised”, “established” and even “overrun” more or less synonymously; likewise Thomson, for fauna in The Naturalisation and its derivatives. George Thomson’s treatment of introduced plants relied on descriptors such as “common on poor, rather sterile land,” “occurring in fields waste places,” “a fugitive garden escape,” etc., which would probably have lent themselves to a formal classification had he or his informants been aware of such categories, or perhaps just willing to apply them.167 Thomson’s “weeds of cultivation” remark (above), almost directly echoed published phyto-anekeitaxa, such as Watson’s 166 Ibid., 82. 167 Thomson, The Naturalisation, (cit. n. 150); e.g., 460; 479; 480. iv 83 “colonists” and Thellung’s “epœkophytes.” An unawareness of Thellung’s thenrecent works, published only in French and German, would be less surprising than unfamiliarity with Watson’s older, simpler, near-colloquial English terminology. Kazimierz Wodzicki’s table of contents offered clues to his anekeitaxonomic conceptions. Several categorical subheadings recur, taxon by taxon: liberations; early spread; present distribution; economic value (or status, or importance); limiting factors; and control. In a few cases, these are truncated or expanded. “Liberations” provides both priority of occupation and assigns responsibility or blame for the introduction. “Early spread” suggests something now often called “invasiveness,” and more subtly, the point up to which decisive actions could or should have been to taken to contain the beasts. “Present distribution” and “Limiting factors” correspond to different pseudometric degrees of naturalization. “Economic” factors and “Control” dealt both with perception and enforcement of a taxon’s social tolerability. Some of his species accounts underline these judgments by discussing “infestations” rather than “populations.” Others are burdened with incommensurable, vague and potentially overlapping terminology: rare, common, numerous, widely (vs. locally) distributed, feral, semiferal, well-acclimatized, established, well-established, naturalized, and even “occupy[ing] all suitable areas.” Taken as a group, these concepts suggest, at iv 84 best, a fuzzy, multi-dimensional status grid rather than a set of well-developed categories.168 Ecology Textbooks Accounts of neobiota appeared in several ecology textbooks, dating from 1895 to 1954.169 As translated (1909), Eugenius Warming’s 1895 Plantesamfund included about half a page devoted to neobiotic plants. “Hordes of European plants … have entered Argentina and have destroyed the indigenous vegetation here and there. On the other hand, American plants introduced into Europe have locally suppressed native species … in the same way, several hundred foreign species have reached New Zealand, where some of them defeat the native vegetation. It is essential that climate and soil shall suit immigrant plants; otherwise, they fail to gain an entrance, even when protected by man.”170 Coulter, Barnes and Cowles’s 1911 Textbook of Botany closed its section on “plant associations” with just under a half page of “The influence of man upon vegetation:” Man is the most destructive of animals … responsible for distributing through the world most of the “weeds” which burden the farmer and throng the roadsides … plants of this sort that inhabit fields and waste places are known as ruderals … The pioneer associations that follow in man’s destructive train, such as the ruderal associations of fallow land or the “fireweed” association of a burned forest tract … often contain xerophytic species … if man leaves such areas to their natural course, there is 168 Wodzicki, Mammals (cit. n. 160). 169 Charles Elton’s pre-1958 books are discussed in chapter two. 170 Eugenius Warming, Œcology of Plants Trans. Percy Groom and Isaac B. Balfour. (Oxford: Clarendon Press 1909), 364. iv 85 succession of associations … culminating finally in the plant association that originally dominated the region”. 171 Victor Shelford’s 1913 Animal Communities in Temperate America included scattered references to introduced biota, particularly in his discussion of “secondary communities.” But he did not always acknowledge them as such (e.g., carp) and conspicuously omitted any mention of then-notorious cases such as house sparrows and starlings. 172 As translated (1937), Richard Hesse’s 1924 Tiergeographie auf œkologischer Grundaige included an eighteen-page, closing chapter titled “The Effect of Man on the Distribution of Animals.” Just over two pages dealt with “Intentional and unintentional transport by man.” Almost the whole subsection was taken up with reciting a litany of introduced species, followed by a paragraph on quarantine measures, and a final thought: “Thus man attempts to restore the balance of nature which he himself has destroyed; despite obvious difficulties, [quarantine] has now become a standard procedure in pest control.173 Allee, Emerson, Park, Park and Schmidt’s 1949 Principles of Animal Ecology offered about seven pages examining the “unwitting [neobiotic] experiments.” They concentrated on ideas such as the apparent “preadaptation” of successfully introduced species, the existence and operation of “biotic 171 John M. Coulter, Charles R. Barnes and Henry C. Cowles A Textbook of Botany for Colleges and Universities (New York: American Book Co. 1911), 946. 172 Victor Shelford, Animal Communities in Temperate America (Chicago: University of Chicago Press 1913). 173 Richard Hesse, Ecological Animal Geography. Trans. W.C. Allee and Karl P. Schmidt. (New York: John Wiley & Sons, 1937) 551-553 iv 86 barriers” to immigration, the role of human-modified habitats, the characteristics of island biotas, and the competition between congeners. To their satisfaction, all the cases they discussed recommended the conclusion that animal communities are supraorganismic entities.174 Eugene Odum’s 1953 Fundamentals of Ecology devoted the last paragraph of a section on wildlife management to “a comment on the introduction of exotics, that is, of species not native to the region.” His pragmatic conclusion: In general, if there is a suitable species native to the region, it is better to concentrate on management of this species than to introduce a substitute. If, on the other hand, the environment has been so changed by man that native forms are unable to survive, introduction of a species adapted to the new environment may be in order.”175 Odum did not elaborate on the procedures for selecting such substitutes. H.G. Andrewartha and L.C. Birch’s 1954 The Distribution and Abundance of Animals did not address neobiota as a discrete class of phenomena. They used brief examples of species introductions in several sections of the book to make or support particular contentions. Rabbits, which might seem likely to interest Australian authors of such a book, were mentioned only in passing; the interaction of Opuntia and its herbivore Cactoblastis received more attention.176 174 W.C. Allee, Alfred E. Emerson, Orlando Park, Thomas Park and Karl P. Schmidt, Principles of Animal Ecology. (Philadelphia: W.B. Saunders, 1949). 175 Eugene Odum, Fundamentals of Ecology (Philadelphia: Saunders 1953), 328. 176 H.G. Andrewartha and L.C. Birch The Distribution and Abundance of Animals (Chicago: University of Chicago Press, 1954), 87, 337, 501. iv 87 Conclusions This chapter demonstrates that neobiota have been of religious, economic and theoretical interest to scientists since the self-conscious beginnings of modern science. European countries, whether aspiring to trans-oceanic empire or convinced that such empire was beyond their grasp, took note of the fact that some plants and animals could be moved from place to place and established anew. Pioneering biogeographers and taxonomists had to reconcile their new learning with Biblical accounts, or show that their empiricism rendered those traditions untenable. The fact that some plants and animals could not only survive, but even thrive in new lands challenged notions of an orderly creation; the corollary that in so thriving they might replace indigenous island biota (which were perhaps created to “fit” their unique conditions) added fuel to that fire. A tradition that I call anekeitaxonomy, comprised of attempts to categorize plant species according to degrees of geographic belonging, took root in the first half of the nineteenth century. Some of its terminology persists in modern invasion biology, particularly as practiced in central Europe. But anekeitaxonomy is unwieldy. It rests on a quasi-legal conception of a right or privilege of prior occupation. Its “was = ought” construction defies geological dynamism and even evolution, but that is reasonable given its pre-Darwinian conception. It is far more interesting that not merely associating species with geographies, but permanently attaching them, survived the “Darwinian Revolution” that set so many other certainties of natural history adrift in a sea of contingency. iv 88 A whole new “ecological” rationalization for asserting the attachment of biota to places to had to be developed after the (first) collapse of creationism. That rationalization would be cobbled together from fragments of ongoing observation and theorizing, alloyed with continued, if diminished respect for Biblical (or ecclesiastical, or perhaps just cultural) tradition, and with persistent if vague notions of wildness, savage nobility, pre-lapsarian perfection, posttechnological nostalgia, and the appalling ravages of “total war.” But ecological textbooks took varying notice of and stances regarding neobiota, and there was no obvious trend among their treatments from 1885 to 1954. iv Chapter 2: The Mutual Admiration of Charles Elton and Aldo Leopold Among scientists, disciplinary identities have certain tribal attributes, one of which is that we of the tribe are the people, and they, outside the tribe are not the people (or at least, not the people who matter). Put more directly, what scientists outside our discipline have to say doesn’t matter, so we cite only those who practice our own tribal science.177 Even in the nineteenth century, before subdisciplines really proliferated, zoologists and botanists attracted to the study of neobiota were not paying much attention to each other. The boundaries only solidified in the twentieth century, and this continued to affect the study of neobiota. But within the uncertain boundaries of its infancy, ecology allowed a Yale forester and an Oxford zoologist to begin a conversation that still echoes today. Aldo Leopold (1887-1948) and Charles Elton (1900-1991) lived in a world where place mattered. Where you were, where you could go, where you spent your time not only demonstrated, but helped determine whom and what you were. That, of course, was hardly new a century ago, and remains the case today. But there is a difference between recognizing that place matters and enforcing it, should the unthinkable “out of place” exception occur. Leopold and Elton were born into two sorts of middle class families, both reaping the benefits of the status quo, and both depending on its persistence. Each devoted his life 177 See, e.g., Thomas Gieryn, Boundaries of Science, IN Sheila Jasanoff, et al, Eds. Handbook of Science and Technology Studies. (Thousand Oaks, CA: Sage Publications, 1995.) 393-443. 90 to understanding some of the processes that maintained it. And when the unthinkable occurred, both volunteered to help enforce it. By the turn of the twentieth century, it was clear to both zoologists and botanists that their objects of study were regionally differentiated, but science was no longer preoccupied with reconciling Genesis and geography. They had natural selection to account for biotic diversity, and uniformitarianism to account, at least in part, for topography. They could map the geographic distribution of life in considerable detail, but lacked strong theories to account for planetary scale physical geography or the manner in which biotic distributions had developed. Describing the diversity of local biotas still substituted for explaining them. Everyone recognized the effects of latitude, altitude, exposure and climate. God may not have placed plants and animals at selected centers of creation, but they lived where they did for very good reasons. At the zoo, or the botanical gardens, or the natural history museum, specimens were labeled with a country or continent or island of origin. These attachments were organic, dependable facts. The new, ecological descriptions assumed that physical geography itself determined biotic distribution. If removed or “disturbed,” they inevitably (unless actively prevented) obeyed geographical dictates by returning, via succession if necessary, to their marks on the world stage. At least at the scale of an individual human observer, biogeographical dynamism was trivial, local and repetitive. The natural world had boundaries, and biota did not merely respect them, it reflected them. 91 Culturally, old outsiders were becoming the new insiders. Despite improving long-distance transportation and communication making it theoretically easier to administer and exploit a global empire, European colonies were asserting or reasserting their independence. As their ties to “home” countries weakened, they became places in themselves (and vice versa). Ex-colonists, now citizens, asserted local identities, and recovered, restored, or invented boundaries between themselves and imperial centers. More places mattered, in and of themselves. By 1900 the boundaries of the contiguous United States had been stable for half a century. Internal exploration and discovery gave way to infill settlement and efforts to manage resource exploitation, and the first American forestry school opened at Yale. Calls to preserve some areas, unsettled and unexploited, had already led to the establishment of several national parks. In cities, zoning laws and regulations were becoming more common. The common wisdom, however diversely expressed, was that everything had its place, and should generally expect to remain in it. Charles S. Elton attended Oxford University and then stayed on for some forty-five years, during which he founded and directed the Bureau of Animal Population (BAP). In the late twentieth century, Elton’s 1958 book The Ecology of Invasions by Animals and Plants was identified as a founding manifesto by a nascent conservation subspecialty calling itself ‘invasion biology.’ Participants and apologists of the field suggested that invasion biology coalesced in Elton’s mind as (e.g.,) a matter of thoughts “first pulled together … in 1957,” presumably 92 either upon mature reflection or in a “eureka!” moment grounded in decades of study.178 But even at that, there is no acknowledged tradition among invasion biologists that the name of their field commemorates Elton’s formulation.179 Elton was the founding editor of the journal Animal Ecology, published nearly 100 scientific papers and several books of his own, but rarely played the public intellectual. He exploded onto the ecological scene and made his reputation with a 1927 synthesis, Animal Ecology. Fifty years later his career was affirmed as a success with the award of a Tyler Prize for Environmental Achievement, “the premier award for environmental science, energy and medicine conferring great benefit upon mankind,” widely touted as “ecology’s Nobel.”180 Elton’s prize noted: “Often considered as the father of the science of animal ecology, his fundamental research has led to a greater understanding of ecological competition, and the organization and disruption of animal communities.”181 How did Charles Elton develop his conclusions regarding neobiota? Did they spring from his brow, Athena-like, wholly formed in Invasions? Is there evidence of a longer process? This chapter sketches an answer to those questions, and others that arose along the way. In particular, the philosophy 178 Baskin, A Plague of Rats and Rubbervines, (cit. n. 6) 8-9; 128 and first edition dust jacket. 179 James T. Carlton to M. Chew, 25 September 2006. 180 Anonymous, Tyler Prize Homepage, accessed online 25 September 2006 at http://www.usc.edu/dept/LAS/tylerprize/index.html. 181 Anonymous, Tyler Prize Laureates (undated) accessed online 25 September 2006 at http://www.usc.edu/dept/LAS/tylerprize/previous.html. 93 Elton espoused in the final chapters of The Ecology of Invasions by Animals and Plants is strikingly reminiscent of Aldo Leopold’s prior, better-known “land ethic.” Elton did cite the American once in the book, but only in another context. Did Leopold influence Elton, or vice-versa, or both? What connected the two men and their ideas? A great many things connected them. Elton’s and Leopold’s attitudes about neobiota arose early in their respective careers, based on particular local cases, and were thus relatively simplistic, perhaps immature judgments. Neither man substantially deviated from his early assessment, and each gradually intensified his rhetoric as his career progressed. The previously unexplored relationship between Elton and Leopold is revealed in their own words, showing the interdependence of their ideas and suggesting that their careers must be understood in mutual context. Two highly accomplished natural historians are shown eliding ecology’s theoretical pitfalls by means of commonsense analogies and metaphors, literary and cultural allusions and (often) elegant prose. In the process, invasion biology’s simple, Eltonian “creation myth” collapses, and is replaced by a more tenable narrative of conceptual inertia and rhetorical evolution. Susan Flader and J. Baird Callicott wrote, “Leopold and Elton…had their own mutual admiration society grounded in appreciation of each other’s ability to conceptualize new advances in ecology and express them in language any 182 thinking individual could grasp.” 94 But just as G.E. Hutchinson’s n-dimensional niche eclipsed Elton’s conception of the niche as an ecological “profession,” a more nuanced approximation of the Elton-Leopold relationship is possible. The documentation is disappointingly thin, and it is tempting to attribute meaning to interstices that might merely signify a poorly preserved record. Nevertheless, enough remains to compose a sketch showing these two acclaimed men discussing matters both profound and ephemeral, while revealing a few very human foibles and faults. Their myths may be diminished as these human dimensions are reinstated, but neither man seems to be in serious danger of losing his legendary status as a result. The State of the Existing Record Aldo Leopold’s life and work have been documented and analyzed at considerable length. He was born into a prosperous Iowa family early in 1887, two generations removed from German immigrants. His father was an outdoorsman and the setting in which they lived was congenial to hunting and nature study. Shy but sedulous, Aldo was a good student. By age fifteen he was keeping extensive records of his bird sightings.183 He followed his outdoor interests, rather than the family furniture business, through his education and throughout his career. His health was far from perfect, but his death was unforeseen, “in the midst of life.” Biographies and biographical commentaries by 182 Susan Flader and J. Baird Callicott, eds. The River of the Mother of God and Other Essays by Aldo Leopold. (Madison: University of Wisconsin Press, 1991), 224. 183 Curt Meine, Aldo Leopold (cit. n. 163), 11, ff. 95 Curt Meine, Susan Flader and others cover most of this territory, and are freely cited here. If there is any shortcoming in that body of work, it is a hagiographic leaning. Leopold is revered, and unapologetic critiques of his work are rare. But the extensive collection of his correspondence and other personal papers at the University of Wisconsin remains a rich source of unrefined ore. Charles Elton remains a little understood, enigmatic figure. His life is documented in summaries, most of which were appreciations written to mark his retirement or death. Intensely “private,” retiring and even diffident, he avoided conferences, meetings and social occasions. He resisted most attempts to celebrate the scientist rather than his science, even excusing himself from traveling to California to collect his gold Tyler medallion and prize money. He defied the entreaties of biographers, including longtime colleagues.184 Would-be Elton biographers including the prolific historian of ecology Frank N. Egerton reported getting “the cold shoulder” from Elton when they requested interviews or access to historic documentation.185 Elton’s retirement was a long one (1967-1991), during which he apparently winnowed his papers ruthlessly in order to maintain a permanent privacy. He compiled much of what they might have revealed into unpublished narratives of his own, which reside in archives at Oxford and Cambridge 184 See, e.g., Peter Crowcroft, Elton’s Ecologists (Chicago: University of Chicago Press, 1991) x-xi. 185 Frank N. Egerton, personal communication, 8 August 2006. David Cox (personal communication, 16 August 2006) reported that (twenty years after Egerton’s experience) Elton was no longer flatly uncooperative, but he insisted on reviewing and editing Cox’s notes of their discussions. 96 Universities. But bits of the Eltonian disjecta membra persist among the papers of his many correspondents and the organizations that funded his work. A side effect of reassembling some of these materials was determining that Elton’s retrospective accounts and the evidence of surviving documents (including his own contemporary notes) sometimes diverge to a degree better accounted for by active revisionism than fading memory. A few that bear on the topics at hand and significantly alter the received Elton legend are described here in detail. Until a more comprehensive comparison is undertaken, accounts of events based wholly or mainly on Elton’s narratives, without strong corroborating sources, must be considered provisional. This alone “unsettles” much existing scholarship, which largely relied on Elton’s accounts. Like Aldo Leopold, Elton is held in high retrospective esteem; but he is less widely known. His writing was more often aimed at a professional audience and less often advocacy-driven than the American’s. He was remembered by his Oxford students and colleagues as a “genius,” and left a lasting impression of deep thoughtfulness.186 Most of his ecological ideas have been superseded or refined by later investigators, and his methods have never provided answers to some of the questions to which he applied them. But, in the words of his colleague Dennis Chitty, “It’s always gratifying to see one’s ideas taken seriously, if only to see them rejected or modified.”187 186 187 Crowcroft, Elton’s Ecologists (cit. n. 184), 127. Dennis Chitty, Do Lemmings Commit Suicide? Beautiful Hypotheses and Ugly Facts (Oxford: University Press, 1996) 191. 97 One striking thread running through appreciations and reminiscences of Elton (and uniting them with those regarding Leopold) is that they do not credit him with any major theoretical contribution to ecology. Rather, they concentrate on his qualities as an outstanding natural historian, writer, mentor and reviewer. Each man also managed to make his academic base of operations an object of pilgrimage for both established and aspiring colleagues. The way they accumulated such authority can only be hinted at here, but some of the ways they applied their influence is made more apparent. Except as narrowly highlighted by his retroactive acclamation as the founder of invasion biology, Elton’s advocacy of natural resource conservation is little known outside Britain. Even there, since he prided himself on wielding behind-the-scenes influence, he is not widely remembered for his contributions. It seems unproductive to ask whether his interest in neobiota drove his more general interest in conservation or vice versa; either way, the two are inseparable. It seems easier to say that conservation inspired Leopold, since introduced species feature explicitly in only a few of his conservation writings. I began this part of the study by focusing on Elton’s writings about neobiota, and moved to Leopold’s as suggestions of a significant relationship between the two began to emerge. A few documents found among Elton’s papers and many more among Leopold’s revealed something of their personal relationship and suggested a great about the development of their shared conservation philosophy. All these stories seemed to merge into a single narrative without much prompting from me, so I present them here as such. 98 First Statements by Leopold: Problems with Trout and Cats The earliest dated statement I have seen by Aldo Leopold mentioning neobiota was a short paper written for an Annual Meeting of the American Fisheries Society (which he had just joined).188 It was late August 1917. Leopold was thirty years old, twelve years out of the Yale forestry school and working for the U.S. Forest Service in Albuquerque New Mexico; his third child (and first daughter) Nina was born that month.189 Leopold did not attend the meeting. “Mixing Trout in Western Waters” was “read by title” into the proceedings along with those of other absentee contributors.190 It was later published in the Society’s Transactions. In the paper, he asserted, “Rainbow, eastern brook, cutthroat, and German brown trout have been indiscriminately mixed with the native black-spotted trout of our southwestern streams.” He followed briefly with a string of anecdotal, admittedly “meagre” observations, things “commonly believed by fishermen,” and “merely indicative impressions.” He concluded that “species spawning at the same time may hybridize,” and “these hybrids are less productive, and thus less desirable than pure stock.” The paper closes with three “rules for stocking practice.” Last among them: “Stocked waters will not be further mixed. Restock with the best 188 John W. Titcomb, “Proceedings of the Forty-Seventh Annual Meeting of the American Fisheries Society, St. Paul, Minnesota, August 29, 30 and 31, 1917.” Transactions of the American Fisheries Society 47 (1918), 51-79. 189 Meine, Aldo Leopold (cit. n. 163) 162. 190 Titcomb, “Proceedings” (cit. n. 188) 77. 191 adapted species, natives always preferred.” 99 He lacked data, and even a testable hypothesis, but that did not deter him from drawing a conclusion and taking a regional, nativist stand. To all appearances, trout introductions simply violated Leopold’s sense of piscigeographical propriety. An undated Leopold typescript headed “Birds and Cats” presumably corresponds to an April 1918 Albuquerque Evening Herald (newspaper) article by that title.192 Opening with a militaristic analogy redolent of the times, Leopold wrote: “When talk[ing] of the pleasant subject of peace, we have also got to deal with the unpleasant subject of war. Likewise when we talk about birds, we have got to deal with cats.” After a few remarks about the long history of cat keeping, he embarked on an anti-feline polemic. Some of his points seem strained. Paraphrasing a familiar Darwinian anecdote, Leopold argued that the abundance of mummified cats known from ancient Egypt suggested that ancient Egyptian birds were scarce, and thus ancient Egyptian fields would have been ravaged by insects—which “might have been at least in part responsible for the periods of crop failures and famines which preceded the downfall of Egyptian civilization.”193 191 Aldo Leopold Papers, 1903-1948, University of Wisconsin, Madison. Archives and Records Management (ALP) 9/25/10-6 17 005,; Aldo Leopold, “Mixing Trout in Western Waters.” Transactions of the American Fisheries Society 47 (1918) 101-102.. This paper was omitted from the “Publications of Aldo Leopold” by Flader & Callicott in The River of the Mother of God (cit. n. 182) (349-370); but when queried (email, 3 February 2006) Flader replied “I am quite familiar with this item and cannot explain why it is not in my bibliography.” 192 Listed in “Publications of Aldo Leopold” Flader & Callicott, The River of the Mother of God, (cit. n. 182) (349-370). 193 In contrast, Helen Macdonald [Falcon, (London: Reaktion, 2006) 48-51] argued that the abundance of mummified falcons found in Egyptian temples and funerary materials boded poorly for the falcon population, not for their prey. 100 He dismissed the value of keeping cats for rodent control, and very roundly but “very conservative[ly]” estimated an annual toll of “250,000,000 birds that we must charge against the cat nuisance.” Furthermore, he wrote, cats consumed foods “which ought to go toward the feeding of chickens or other producing animals,” and they were “known to be a serious factor in the dissemination of contagious disease.” He predicted: “The time will come when it will be illegal to keep a cat without a license just as we now license a dog, and when all unlicensed animals will be disposed of by public authorities.”194 Ten years later, Leopold wrote to Wilson Bulletin editor T.C. Stevens [sic] at Morningside College in Iowa, regarding a recent Bulletin article.195 Leopold was concerned about the possible need for two discrete sets of “control measures” for housecats, which would have different winter habits in cold and mild regions. He inquired whether Jean Linsdale, the author of the article, had “observed any cats which seemed to be truly wild in the sense of not making their headquarters at any human habitation? Evidently there is a zone terminating somewhere in Iowa where the cats can stay outdoors the year long.”196 On the 194 Aldo Leopold, “Birds and Cats” ALP 9/25/10-6 017 008. To provide some indicator of the modern picture, Google web searches (3 February 2006) revealed that the term “dog license” occurred almost eighteen times more frequently than “cat license” (277,000 vs. 15,600 hits among searchable items) and that Albuquerque was among the jurisdictions requiring cats to be licensed. At mostly non-U.S. sites (“licence”) the ratio was higher, with 63 times as many “dog” hits. Finally, Monty Python’s “fish licence[-ense]” sketch, which mentions both dogs and cats was excluded from the foregoing, but scored about a thousand hits. 195 His name was actually T.C. Stephens. 196 Aldo Leopold to T.C. Stephens 17 October 1928. ALP 9/25/10-4 004 013. 101 specified page, Linsdale had actually stated, “Destruction of some species of birds has been increased with human settlement of the area. Two important enemies (cat and dog) were brought in.”197 Leopold’s concerns apparently did not extend to dogs, which often appeared in his own family photos. Flader and Callicott identified Leopold’s unpublished 1923 essay “Some Fundamentals of Conservation in the Southwest” as his “first sustained effort to express a rationale for conservation beyond narrow utilitarian criteria.”198 After indicting current land management practices on a historical basis for several pages, he turned to “conservation as a moral issue.” Leopold invoked the ideas of Russian mystic philosopher Peter Ouspensky, but perhaps he could just as easily have extrapolated from the elaborate, superorganismic, successional conceptions of ecologists Victor Shelford (1913) and Frederick Clements (1916).199 Thus Leopold first rationalized the ethical dimension of his mostly practical conservation ideals: [Ouspensky] states that it is at least not impossible to regard the earth’s parts—soil, mountains, rivers, atmosphere, etc.—as organs, or parts of organs, of a coordinated whole, each part with a definite function. And, if we could see this whole, as a whole, through a great period of time, we might perceive not only organs with coordinated functions, but possibly also that process of consumption and replacement which in biology we 197 Jean Linsdale, "Some Environmental Relations of the Birds in the Missouri River Region." Wilson Bulletin 40 (1928): 157-177. 198 199 Flader & Callicott, The River of the Mother of God, (cit. n. 182) 13. E.g., Victor Shelford [Animal Communities in Temperate America, (cit. n. 172) 35] wrote: “Communities are systems of correlated working parts. Changes are going on all the time … much like the rhythm in our own bodies related to day and night;” Clements [Plant Succession (Washington D.C.: Carnegie Institution, 1916), 30] invoked “a complex [ecological] organism, possessing functions and structures, and passing through a cycle of development similar to that of a plant." 102 call the metabolism, or growth. In such a case we would have all the visible attributes of a living thing, which we do not now realize to be such because it is too big, and its life processes too slow. And there would also follow that invisible attribute– a soul, or consciousness– which not only Ouspensky, but many philosophers of all ages, ascribe to all living things and aggregations thereof, including the ‘dead’ earth.”200 Early Eltonia: Sibling Authority, Arctic Blowflies, and a War in the Parks Charles Elton’s father Oliver was a well-known and respected literary critic, a Professor of English whose circle of friends included the Huxleys and other influential, intellectual British families. Meanwhile, his mother Letitia wrote for popular magazines, and in 1906 dedicated “a tiny book called The Story of Sir Francis Drake” to Charles, her youngest (then five-year-old) son.201 Elton’s oldest brother, Geoffrey, seven years senior, was a critically important contributor to his young sibling’s career choice. As Charles later put it, “without the inspiration of Geoffrey’s love of natural history, I might not ever have become an ecologist.”202 Geoffrey died in his early thirties, but his influence lasted Charles a lifetime. Even well into his (active) eighties, Charles often reflected on his adventures and nature study with Geoffrey, and related them in detail.203 We know relatively little about Geoffrey, mostly by way of Charles’s reminiscences and a few surviving letters. He attended Cambridge and worked 200 Flader & Callicott, The River of the Mother of God, (cit. n. 182) 95. 201 Charles S. Elton Papers, Oxford University Bodleian Library MS.eng.c.3326. (CSEP) A33:11; Mrs. Oliver Elton, The Story of Sir Francis Drake (London: T.C. & E.C. Jack, 1906). 202 CSEP A34: 14. 203 Telephone conversation with David Cox, 16 August 2006. 103 for a time in a fish hatchery, where he became interested in curing fish diseases.204 He drilled with the Army during World War I, but apparently saw no action.205 Ultimately, he taught in “a co-educational school” and drafted a textbook for English teachers.206 He underwent a surgery on his neck that physically weakened him.207 But his letters reveal a strong personality. Geoffrey was a stickler for hypothetico-deductive approaches to natural history, writing to Charles at Oxford: “Before you cut up a new animal, say a crayfish, submit your idea of what would make a good crayfish, where its liver ought to be … etc. Then see how the real crayfish’s effort compares with yours. But you must invent a crayfish before you have seen a single hint of what the real one is like: this is absolutely essential.” On a livelier note, in the same letter, “I believe it’s a good idea, if you’ve been doing a lot of stuffy head-work and want an interval, to get several jars of animals & simply watch them for half an hour or more and see what they’re doing & use your imagination. You don’t need to think about them at all, in fact the less you think about them the more ideas you’re likely to have, and you’ll want to forget all about old theories & start with a perfectly fresh and open mind.” 208 204 C.S. Elton to G.Y. Elton 14 April 1913. CSEP A19; G. Elton to C. Elton 3 November 1920. CSEP A21. 205 G.Y. Elton to C.S Elton 26 June 1917. CSEP A20. Charles himself trained with the Signal Corps, but was never mobilized. 206 C.S. Elton, “The Death of Geoffrey Elton.” Unpublished manuscript. CSEP A34. 207 C.S. Elton, “Small Adventures.” Unpublished manuscript. CSEP A34: 21. 208 G.Y. Elton to C.S. Elton 3 November 1920. CSEP A33. 104 Geoffrey also clearly valued originality: “I didn’t quite follow all you said about what your program was, but it seemed rather as if you were simply working out things that had been very thoroughly worked out before … if you could specialise in some subject very persistently, and … get to know more than anyone else in the world about it, I believe that you would find it a frightfully useful weapon—not as your main occupation, I mean, but as a side-line (to make yourself necessary to people).” Finally, he insisted, “I believe the only way to understand animals is to study them for a useful purpose. They all grow for practical purposes and study each other for definite important ends, and you can’t get into the right atmosphere if you only theorise about them. It isn’t urgent enough, you can’t do any work well unless it’s urgent, at least I can’t.”209 Charles appears to have unquestioningly deferred to Geoffrey; Geoffrey certainly assumed an air of authority over Charles.210 Peter Crowcroft and several of Elton’s eulogists were aware that Elton privately dedicated the BAP as “a dynamic memorial” to Geoffrey, whose 1927 death caused Charles “great and lasting sadness.”211 The BAP persisted, and sometimes prospered, until Elton’s 209 G.Y. Elton to C.S. Elton “Wednesday night” (undated, likely late 1920) CSEP A21. 210 Charles was in awe of his father Oliver, as well. It might not be amiss to bear in mind that whatever authority Elton later wielded, it was likely colored by his boyhood role as “Elton minor.” 211 Crowcroft, Elton’s Ecologists, (cit. n. 184), 8; Richard Southwood and J. R. Clarke, “Charles Sutherland Elton. 29 March 1900 - 1 May 1991,” Biographical Memoirs of Fellows of the Royal Society 1999, 132-140. 212 retirement. 105 His unhappiness at its dissolution by his Oxford successors was necessarily aggravated by what he saw as a desecration of his brother’s cenotaph. Elton first published evidence of his interest in neobiota with his 1925 paper “The Dispersal of Insects to Spitsbergen.” There he wrote, “The sealing ships which visit Spitsbergen carry with them a small insect fauna … flies breed in the remains of herrings which are carried as cargo in the winter months to the Roman Catholic countries of southern Europe… It is thus apparent that the dispersal of insects to Spitsbergen is partly dependent on religious movements. So far, none of these species seem to have established themselves…except blowflies.”213 But a more domestic case was already on his mind. Late the same year, Elton wrote to his mother, “The [red] squirrels are … conducting a very hard campaign: they were defeated in Wadham gardens about 1921, and driven out of the University Parks about 1924, and finally left Magdalen Park this year. The advance guard of grey squirrels has just reached Bagley wood, but Shotover and 212 Crowcroft (cit. n. 184), Southwood and Clarke (cit. n. 211) and David L. Cox, [Charles Elton and the Emergence of Modern Ecology. (Dissertation, Department of Biology, Washington University, Saint Louis, MO. 1979)] all wrote substantially about Elton and the BAP. 213 Charles S. Elton, “The Dispersal of Insects to Spitsbergen,” Transactions of the Entomological Society of London 73: (1925) 89-299. Unlooked-for ecological ramifications of religious practice would attract his attention later, and were included in his 1958 book. 106 Blenheim park are still pure red. We have a big six-inch map of the district, and I put various coloured pins in, as people bring information in. It is quite warlike.”214 Reminiscent of Charles Darwin (see chapter one), Charles Elton imputed the nationalist and martial characteristics of his own culture and its recent Great War (e.g., border defense, military organization, and strategic activity) to rodent populations without reflecting (for the record, at least) on the pitfalls of doing so. Some general insight might attend Elton’s then-recent experience of drilling with the Army in the waning days of the war; military training and discipline are meant, among other things, to promote and reinforce such a worldview. Further insight into the squirrel “war” comes to us from Elton’s narrative of his own early childhood. At age four, he was presented with his first book, Squirrel Nutkin, in which (red, English) squirrels captured and delivered other small mammals as tribute to an owl, in return for the right to safely gather nuts.215 It would be strange indeed if this anthropomorphic, reactionary and earnestly prescriptive story either founded or fed Elton’s conception of animals. But late in life he raised such a possibility by writing that Beatrix Potter’s writing “was and still seems to me without a blemish of any kind.” In the same paragraph he also disclosed, without apparent regret, “I was fed on a great deal of War.” 216 214 C.S. Elton to Letitia Elton, 11 November 1925 CSEP A15. British Ordnance Survey maps were scaled at six inches to one mile. 215 Southwood and Clarke, “Charles Sutherland Elton,” (cit. n. 211). 216 C.S. Elton, “Small Adventures.” CSEP A33. 107 How Charles Elton Defined a Field of Study (And How He Didn’t) When Charles matriculated at Oxford, Julian Huxley became his tutor and second de facto mentor. While Geoffrey’s approach seems to have governed his little brother’s approach to field studies and natural history, Huxley supervised his readings, and looked after his professional advancement. Of most readily apparent significance, Huxley edited a series of “Text-books in Animal Biology.” Recognizing his protégé’s potential, Huxley assigned Elton the task that catapulted him to sudden senior status: authoring a summary and synthesis of the emerging topic of animal ecology. Plainly titled Animal Ecology, Elton’s book assembled a set of ideas that had been accumulating for decades into an intelligible-seeming whole. He “developed a set of ‘organizing principles’ for the study of animal ecology, which set the agenda for much subsequent research and theory formation.”217 In thus codifying a new field of endeavor, Animal Ecology remains Elton’s most enduringly significant work. Over time, this youthful (his own and the field’s) effort was eclipsed in some of its fundamentals, and many of its details, but for some thirty years it remained a standard reference.218 But even at the outset Animal Ecology had shortcomings. Elton’s stinginess in crediting earlier investigators was noticeable. And he sometimes 217 Kurt Jax, History of Ecology” IN Encyclopedia of Life Sciences. (London: MacMillan/Nature Publishing Group, 2001), Vol. 9: 55-61. 218 Historians including Robert P. McIntosh [The Background of Ecology: Concept and Theory, (Cambridge: University Press, 1985)] have described the rise, apotheosis and supplanting of Elton’s various ecological ideas, which is beyond the scope of this paper. 108 failed to acknowledge the existence of parallel or alternative views, suggesting better basic theoretical agreement than practitioners had actually achieved. One early reviewer noted in Science that conceptions of the niche other than Elton’s own had received short shrift.219 In an oft-repeated anecdote, Elton later claimed that he wrote Animal Ecology in eighty-five days.220 But correspondence with his well-published (and not-to-be trifled with) father makes it clear that the 1927 book was already seriously contemplated by late 1924. In Charles’s words, “Dear Daddy … you will scream with laughter at hearing that I am going to write a text-book of ecology. Laugh on. Don’t mind me.”221 Indeed, Charles was under contract to produce Animal Ecology by early 1925.222 He reported significant progress (“I have written a great deal of it”) almost exactly a year later.223 If, in any meaningful sense, Elton wrote the book in eighty-five days, they followed either a heroic bout of procrastination, or careful, extended preparation. Charles dedicated Animal Ecology to Geoffrey, who died suddenly of a stomach hemorrhage in the year of its publication. It is not clear whether he lived to see the book and its dedication in print. 219 Taylor, Walter P. 1928. “Review of Animal Ecology by Charles Elton.” Science 68 (1928) 455-456. 220 Southwood and Clarke, “Charles Sutherland Elton,” (cit. n. 211), 143. 221 C.S. Elton to Oliver Elton, 16 November 1924. CSEP A14. 222 C.S. Elton to Oliver Elton, 14 March 1925. CSEP A14. 223 C.S. Elton to Oliver Elton, 7 March 1926. CSEP A14. 109 For Animal Ecology, Elton formulated a basic view of neobiota that he never abandoned. He wrote: “Many of the most striking cases of sudden increase in animals occur when a species is introduced into a country strange to it, in which it does not at first fit harmoniously, often with disastrous results to itself or to mankind.”224 But he mentioned very few specific cases, and used the term “invasion” for population irruptions that did not obviously involve human agency. When discussing migrations, Elton suggested, “It seems highly probable, although difficult in our present state of knowledge to prove conclusively, that many animals migrate on a large scale in order to get away from a particular place rather than to go towards anywhere in particular.”225 He did not, however, expand the conception and suggest that populations established as a result of human activity exemplified successful, if unorthodox, migratory “escapes.” Elton’s Contributions to Wells, Huxley and Wells’ The Science of Life In 1925, Julian Huxley also agreed to work with H.G. Wells (and his son G.P. Wells) to produce a comprehensive biology book for popular consumption. Originally published in serial form, it was later collected into multivolume and single volume books. Likely some time after (and conceivably contingent upon) the completion of Elton’s Animal Ecology, Huxley drafted an ecology chapter for 224 C.S. Elton, Animal Ecology (London: Sidgwick & Jackson, 1927) 111. 225 Ibid., 156. 110 The Science of Life, enlisting Elton to “scrutinize” it. Presumably the elder Wells rewrote this chapter, but it is worth a short dissection.226 “The Science of Ecology” garnered fifty pages in The Science of Life, just over three percent of the total text. Its seven subsections were provocatively titled “Ecology is Biological Economics;” “The Chemical Wheel of Life;” “The Parallelism and Variety of Life-communities;” “The Growth and Development of Life-communities;” “The Grading of Life-communities;” “Food-chains and Parasite-chains;” [and] “Storms of Breeding and Death.”227 It took a deterministic view of community succession.228 Arguing further that succession recapitulates evolution, “The Science of Ecology” explained the persistence of “primitive types” by the need for more evolved species to rely on their forbears to continuously create the conditions necessary for advanced life (e.g., by converting rocks into soils).229 Neither of these ideas reconciled easily with Elton’s usual explanations. In The Science of Life, neobiota remained undiscussed until the last “Ecology” subsection. There, a few notorious cases were listed, but (as in Animal Ecology) not strongly categorized as abnormal or different from other kinds of population dynamics. The piece wound down by relating Elton’s Animal Ecology anecdote about the cats of Berlenga Island who eradicated their own 226 Peder Anker, Imperial Ecology: Environmental Order in the British Empire, 1895-1945 (Cambridge, MA: Harvard University Press 2001) 112. 227 H.G. Wells, Julian S. Huxley, and G.P. Wells. The Science of Life. (New York: Literary Guild, 1934) 961. 228 Ibid., 973 229 Ibid., 980 230 food supply, and similar Malthusian horrors. 111 Then came the finale: “Unrestrained breeding, for man and animals alike, whether they are mice, lemmings, locusts, Italians, Hindoos, or Chinamen, is biologically a thoroughly evil thing.”231 The next chapter, “Life Under Control,” had a subsection titled “Pests and their Biological Control.” Here Elton’s influence seems more apparent, and several of his later “favorite” cases studies were mentioned.232 Whether he brought these cases to Huxley’s attention or vice-versa is unclear. Another Little Elton Book about Ecology Elton’s second book, Animal Ecology and Evolution, contained “the substance of three lectures delivered in the University of London in the autumn of 1929, under the title of ‘The Future of Animal Ecology.’”233 Again Elton collected ideas from others, acknowledged them in the preface, and wove them into his own expanding concept. Borrowed ideas included habitat selection by animals (J.C. Farthing), the evolutionary implication of irregular migrations (C.B. Williams) and relative optimum densities of humans and animals (A.M. Carr-Saunders).234 230 Elton, Animal Ecology (cit. n. 224) 114; Wells, Huxley & Wells, (cit. n. 227) 1010-1011 231 Ibid., 1011 232 Ibid., 1016-1032 233 C.S. Elton, Animal Ecology and Evolution (Oxford: Clarendon,1930) 6. 234 According to Jack Morrell [Science at Oxford 1914-1939 (Oxford: Clarendon,1997) 292] Carr-Saunders’ book The Population Problem (Oxford: Clarendon,1922) was an early (and neoMalthusian) influence on Elton’s thinking. Carr-Saunders’ concept of societies adjusting themselves toward an “optimum density of population” may be what Elton had in mind in The Ecology of Invasions (92) when he wrote of a time “when numbers of human beings were regulated by customs, often harsh enough, but meeting the end desired.” 112 And here, for the first time (at least under his own byline) he cited George M. Thomson’s comprehensive 1922 treatment, The Naturalisation of Animals and Plants in New Zealand.235 Among contemporaneous reactions, an anonymous notice in The Quarterly Review of Biology stated noncommittally, “This little book merits the careful consideration of students of biology.”236 But marking Elton’s retirement in 1968, Sir Alister Hardy recalled the slim volume in glowing terms: “Few books on Zoology have delighted me more than did this when it first appeared.” Among those delights, he related Elton’s empirically-informed take on habitat selection: “’Now what we know of the real, as opposed to the textbook, behaviour of wild animals establishes the fact that they do not remain still to be selected. In periods of stress it is…common for animals to change their habits, and this usually involves migration.’”237 Focusing on neobiota, Elton wrote, “With the tremendous radiating power of the world’s transport, the introduction of alien animals is now almost a daily occurrence, and the subsequent increase of some of those species as pests has made the practical control of animal numbers a subject of paramount and acute importance in most countries of the world.”238 Nevertheless, he cautioned, “‘The 235 Elton, Animal Ecology and Evolution (cit. n. 233) 85. 236 Anonymous, “Brief Notices,” The Quarterly Review of Biology 6 (1931) 223. 237 Alister Hardy, “Charles Elton's Influence on Ecology.” Journal of Animal Ecology 37 (1968), 3-8. 238 Elton, Animal Ecology and Evolution (cit. n. 233), 11. 239 balance of nature’ does not exist, and perhaps never has existed.” 113 Elton reiterated the term “invasions” as pertaining to population irruptions that “bear the same stamp of apparently complete pointlessness.”240 Perhaps for the first time, he used the term “counter-pest” to describe the purposeful introduction of one species to limit the activities of another.241 In a moment of reflection reminiscent of Geoffrey’s anatomical admonitions, he wrote, “Your ideas upon the origin of what you are studying unconsciously influence your outlook on it, and you cannot really understand the significance of a thing without having some conception of its past history.”242 In summarizing, he waxed philosophical, and perhaps revealed the effects of a childhood steeped in intellectualism, arts and literature. “The real life of animals is therefore a compound of many things: fixed and predetermined limits impressed by the environment; the relations of the sexes; the survival of things that are useful; a certain free will in the matter of choosing between good and evil surroundings, accompanied by a great deal of movement; a fairly large amount of pure chance; and sometimes a growing stock of new ideas born out of contact with new situations– Predetermination, Sex, Materialism, Free Will, Destiny, Originality and Tradition. The study of limiting factors in the surroundings of animals enables us to define the stage and the scenery in the midst of which the act will be played. But the actors, starting though they do with fixed instructions, and fixed limits of time, are still partly responsible for making the play a success or a disaster.”243 239 Ibid., 17. The “balance of nature” is one “commonsense” proto-ecological idea whose popular persistence turned the tables on Elton’s attempts to rhetorically establish ecological theory. 240 Ibid., 45. 241 Ibid., 69. 242 Ibid. 243 Ibid., 90-91. 114 Their Paths Cross: The Matamek Conference on Biological Cycles Aldo Leopold left the U.S. Forest Service in 1928, and began a proposed game survey under the auspices of the Sporting Arms and Ammunition Manufacturers’ Institute [SAAMI].244 Shortly thereafter, Charles Elton secured, then resigned, a minor but permanent position at Oxford, and was “living on grants.”245 The 1931 Matamek Conference on Biological Cycles brought them together. Yale Geographer Ellsworth Huntington summarized this meeting’s purpose: “The Conference was called by Mr. Copley Amory, of Boston, and all of its members were his guests. For years Mr. Amory has been interested in the fact that sometimes the fish, bird, and animal populations of [the North Shore of the Gulf of St. Lawrence] are very abundant, and sometimes, for no apparent reason, very scarce. He has seen years when codfish were extremely abundant, and other years when practically no fish were caught. Salmon and all the other animal populations vary in the same inexplicable way. Therefore, at his summer home on the Matamek [River], Mr. Amory brought together about thirty scientists and Canadian federal and provincial officials to consider the problem of fluctuations in wild life.”246 Elton described it more bluntly: “[T]he idea of it arose 244 Meine, Aldo Leopold (cit. n. 163), 256 ff. 245 Anker, Imperial Ecology (cit. n. 226), 107. 246 Ellsworth Huntington, “The Matamek Conference on Biological Cycles, 1931.” Science 74 (1931) 229-235. 115 from Mr. Amory’s preoccupation with the fluctuations in wild-life resources along the North Shore.”247 Amory appointed Charles to be the Secretary of the conference upon the strength of a referral by a mutual acquaintance at the Hudson’s Bay Company. 248 Elton probably became aware of Leopold, at least by name, during the conference planning. Curt Meine speculated that plant ecologist Frederick Clements might have introduced Leopold to Elton’s work 1930, but that it was “hard to say when Leopold first became aware of Elton. He had likely read Animal Ecology by early 1931, but Elton’s ideas had not yet permeated his approach to wildlife management.”249 Whether either man anticipated their mutual introduction with particular keenness is unrecorded. But it is clear that Leopold almost missed the meeting. In a letter dated 1 February 1931, Alfred Gross of Bowdoin College wrote: My dear Leopold:— I have just received a letter from Mr. Amory in which he states that you have never accepted the invitation to the conference. Therefore I am taking it for granted that it will not be possible for you to attend the program at Matamek next July. If I do not hear from you within the next few days I will suggest another name to Mr. Amory. He says all have accepted the invitation to attend excepting you. I know that you are very busy and porbably [sic] it is impossible for you to get away for the time required for such a trip. 247 C.S. Elton, Voles, Mice and Lemmings: Problems in Population Dynamics. (Oxford: Clarendon),184. 248 249 Southwood and Clarke, “Charles Sutherland Elton,” (cit. n. 211) 138. Meine, Aldo Leopold, (cit. n. 163) 564 n.57; 565 n.57. Meine, among many others, erroneously referred to Elton as “Professor of Zoology.” 116 Handwritten by Leopold across the bottom of the page in urgent telegramese was his apparent reply: “Your letter February one is first notice of invitation to conference I have received stop if not too late I wish to accept stop letter Leopold.”250 Leopold’s archive includes several items related to the Conference: an original, but undated conference invitation signed by Copley Amory and addressed identically to the letter from Gross; a one-page “Abstract of Possible Contributions to Discussion at Biological Conference from Charles Elton;” and his own handwritten notes on Elton’s 24 July introductory talk.251 Once installed at Matamek, Leopold presented “Grouse Cycles and Conservation,” discussing the observed fluxes in several small game populations.252 Elton addressed the conference twice. Summarizing himself, he later wrote, “Elton read an introductory paper on Fluctuations in wild life. He said that practical policies of conservation or production are greatly dependent upon proper knowledge of the factors controlling numbers of wild animals.” 253 Here, perhaps for the first time outside an Oxford lecture hall, he used the seemingly uncontrolled, ongoing expansion of American muskrat populations in Europe as a case study. The muskrat story recurred, with updates, in his later writings. The 250 Alfred Gross to A. Leopold, 1 Feb 1931. ALP 9/25/10-2 005 002. 251 Manuscript headed “Charles Elton. July 24.” ALP 9/25/10-2 005 002. 252 C.S. Elton, Ed. Matamek Conference on Biological Cycles, Labrador, 1931: Abstract of Papers and Discussions. (Canadian Labrador: Matamek Factory,1933) 18-19. 253 Ibid., 15. Leopold’s notes agree with Elton’s official summary of his own statements. 117 second paper, titled “Cycles in the Fur-trade of Canada” was aimed to correspond more precisely with Copley Amory’s “preoccupation.” In a letter written shortly before the conference ended, Leopold called it “the best thing of its kind that I have ever attended.”254 Elton related that (in his experience) it was the only conference at which “the chairman of a session walked in and threw two salmon on the table, announcing with pride that he had just caught them.”255 In 1942, he remembered it as “a remarkable conference, buoyed by Mr. Amory’s personality and vision,” giving “powerful support to a number of ecological projects that were staggering through the trough of the Depression in the human trade cycle.”256 But later, at least, Elton was retrospectively unimpressed with Matamek. He told David Cox “There were some interesting people there, but nothing much of significance happened.” Cox concluded “Elton’s present appraisal … needs to be placed in perspective. Throughout his career Elton has routinely avoided scientific conferences whenever possible. He has often relied on the credible excuse of deafness in his left ear so that he might duck such meetings without giving undue offense. In fact he has rather strong feelings about what he calls ‘conference men’ and the organized farces that fill their time.”257 254 Meine, Aldo Leopold (cit. n. 163), 284. 255 Crowcroft, Elton’s Ecologists (cit. n. 184), 11. 256 Elton, Voles, Mice and Lemmings (cit. n. 247), 185. 257 Cox, Charles Elton and the Emergence of Modern Ecology (cit. n. 212),158. 118 Why a Meeting-hater Risked Two Week’s Seasickness to Attend One Cox believed that Elton attended Matamek in the hope of making contacts that could help him secure research funding, but did not offer any specific evidence. 258 It is now possible to say that by the beginning of 1931, Elton had drafted a narrative and budget proposal outlining a “Research Bureau in Oxford University” listing Copley Amory among his references, and specifically targeting the New York Zoological Society (“NYZS,” below).259 The Society’s archival copy of the proposal is dated 12 January 1931, six months in advance of the Matamek conference; this recommends the conclusion that Elton’s presentations there amounted to a “job talk.”260 Perhaps, just this once, the prospect of funding his proposed research overcame Elton’s shaky sea legs, and his distaste for “organized farce.” At Matamek, Elton favorably impressed Reid Blair, NYZS representative and Director of its New York Zoological Park.261 Cox concluded that afterward, Elton met with Blair and Madison Grant, the Society’s president, and “hammered out an agreement whereby the society began immediate direct support of his research.”262 The details of all this are fuzzy. I have not determined when and where (even whether) this meeting of three minds took place. In 1931 Madison 258 Ibid. 259 C.S. Elton, three memoranda, January 1931. Bronx Zoo Archives. 260 Ibid. 261 Crowcroft, Elton’s Ecologists, (cit. n. 184) 11-12. 262 Cox, Charles Elton and the Emergence of Modern Ecology (cit. n. 212),159160. 119 Grant was approaching the end of a long and distinguished career of nature conservation and advocacy. As a member of the Matamek Conference’s “Permanent Committee” he was expected to attend, but according to Crowcroft, “was in poor health and sent Reid Blair as his representative.”263 If Elton met with Grant on Blair’s recommendation it was necessarily after Matamek and before his 25 January 1932 announcement, establishing the Bureau of Animal Population (BAP) at Oxford “on a temporary and very insecure basis.”264 Crowcroft reported, “the name of the new institute was decided” during a nauseating return voyage to Britain, suggesting that Elton was assured of funding before he departed from North America.265 In any case, in retrospect, Elton could declare that this three-year commitment of funds from New York “kept me alive.”266 Elton’s Brush with the “Patrician Racist” Whether Grant and Elton met and what they discussed might be as pivotal an event as the Society’s funding commitment. In addition to being credited with pioneering conservation triumphs like saving the American bison and California redwoods from extinction, Madison Grant has also been called “intellectually the most important nativist in recent American history.” 267 He authored and edited 263 Crowcroft, Elton’s Ecologists (cit. n. 184), 11. 264 Ibid., 13. 265 Ibid., 12. 266 Cox, Charles Elton and the Emergence of Modern Ecology (cit. n. 212), 167. 267 John Higham, Strangers in the Land: Patterns of American Nativism 18601925. (New York: Atheneum, 1963), 155-157. 120 several “scientific” racist polemics, most notably The Passing of the Great Race (1916) and The Alien in our Midst (1930).268 The juxtaposition at this critical moment of Madison Grant’s rationalized chauvinism and Elton’s ecological nativism is intriguing. It may, of course, be purely coincidental. What Elton knew or thought of Grant’s racist views is unknown, and while Grant may have refined them to an unusual degree and stated them more vehemently and publicly than most, he was hardly unique at the time in holding such beliefs. If Elton went to Matamek expecting to address Grant, his topic of American muskrats invading Europe might have been calculated specifically to harmonize with nativist concerns. It didn’t really speak to cycles as such. Either way, “aliens” made prominent appearances in Elton’s writings shortly after the NYZS became his major patron. Further research on this point is called for, but may be very difficult. According to Madison Grant’s biographer Jonathan Spiro, Grant’s embarrassed family and associates later deliberately destroyed most of the primary source material left by the “patrician racist,” (coincidentally, much as Elton destroyed his own papers). Even the Wildlife Conservation Society (successor to the NYZS), which literally owes its existence to Grant’s efforts, no longer openly acknowledges his contribution to their work.269 268 Jonathan P. Spiro, Patrician Racist: The Evolution of Madison Grant. Ph.D. diss., (Berkeley: University of California, 2000). 269 Wildlife Conservation Society, “Our Mission,” http://www.wcs.org/swour_mission/sw_wcshistory. accessed 23 July, 2006. 121 Beginning a Collaborative Competition Within two weeks of the Matamek conference, Leopold, a bit obsequiously and perhaps on thin pretense, initiated a personal correspondence with Elton. “On the chance that it might be of some service to you, I shall send you from time to time stray bits of information, much of which you will doubtless already have seen, bearing on your Canadian project.” He followed with an anecdote regarding lemmings extracted from a then-new paper about snow geese by George M. Sutton.270 Thus began an exchange that continued at irregular intervals until Leopold’s death. Leopold Echoes Madison Grant and Writes Elton into Game Management At the end of 1931 or early in 1932, a press release appeared bearing a statement attributed to Leopold, and reported in the Journal of Forestry under the title “Game System Deplored as a Melting Pot.” It read, in part: ‘A melting pot that has failed to work,’ is the way Aldo Leopold, wild life investigator of the [SAAMI], senior member of the Society [of American Foresters] and author of the Game Survey of the North Central States, recently described modern game propagation, in a statement made public by the American Game Association. ( … ) Mr. Leopold believes that game restoration agencies have relied too heavily on the makeshift method of substituting foreign birds and species for native game, because of the ease with which these can be raised artificially or imported for restocking from year to year. ‘Thoughtless importation of Mexican quail, which diluted the hardy northern bobwhite blood in Massachusetts to an almost fatal point, is a constant threat against quail future in the rigorous climate to which native birds have become suited. Adding to game variety is commendable and in many sections highly successful, but it must be remembered that native game is still the most popular, that it is truly adapted to its former ranges, that it is fast being forced out by neglect, and that game propagation by 270 A. Leopold to C.S. Elton, 12 August 1931. ALP 9/25/10-2 005 002. 122 way of the incubator and the distributing truck every year is at best a stopgap.’271 The idea that southern races were not only inferior to northern ones but would “dilute [their] blood” can be read charitably as a poorly worded reference to genetic adaptation of animal populations to their local habitat conditions. But it was also, in a nutshell, the argument that Madison Grant made about the dilution of supposedly superior Nordic human stock by southern types in The Passing of the Great Race. The Yale-educated Leopold grew up speaking German and came from mostly northern European stock.272 He could have been unaware of “scientific racism” in 1932, but that seems doubtful. It is evident that he did not practice it at the most personal level. Curt Meine concluded “the Leopold family’s attitude toward racial matters was one of naive indifference tinged by Saxon insularity,” and that working in New Mexico, Aldo himself “had become accustomed to dealing with the full range of Navajos, Apaches, Hispanics, Mexicans, Texans, cowboys of all colors and lumbermen from all corners of the country. He had not anticipated, however, falling in love with [and marrying Estella Bergere,] a Spanish-Italian Catholic.”273 The record is equivocal, but there is no particular reason to suppose that Leopold’s attitude diverged markedly from the spirit of the age. 271 Anonymous, “Game System Deplored as a Melting Pot” Journal of Forestry 30 (1932) 226-227. 272 Meine, Aldo Leopold (cit. n. 163), 15. 273 Ibid.,115. 123 Whenever he first read Animal Ecology, after Matamek Leopold repeatedly made it clear that he was a devotee of Eltonian ideas. His 1933 technical magnum opus, Game Management, included the rarely encountered advice to lay aside the text in hand and pick up another: “If the reader has never read a good text on ecology (such as Elton, 1927 [Animal Ecology]) he may here pause to do so.”274 Leopold summarized the philosophy of his own book by the statement, “Game management consists largely of ‘spotting’ the limiting factor and controlling it. It also equally consists, however, in knowing when to stop, and what other factor next to turn to.”275 The remainder of the book consisted of devilish details. Among them were some related to neobiota, laid out as a sort of continuum: six anecdotal and somewhat fuzzy categories of transplantation failure and success. Employing a confusing idiom that substituted “plant” for “transplant,” he distinguished among “dispersal failure” in which “the planted stock immediately disperses and disappears without breeding;” “straggling failure,” where “the planted stock breeds for one or two years … then persists as non-breeding adults which gradually disappear;” “colony survival”, when “stock persists as a small breeding colony but does not spread … from the viewpoint of producing a shootable game, it is a type of failure.” Successes include “artificial establishment” which requires ongoing intervention like additional stocking to 274 Aldo Leopold, Game Management, (New York, Scribner’s, 1933), 39. This is not as drastic as it sounds; Animal Ecology is a far slimmer volume than Game Management. 275 Ibid., 39. 124 maintain populations; “recessive establishment” where “the stock breeds and spreads … but experiences a subsequent partial decline to a lower level.” and finally “establishment” in which “the stock promptly and vigorously breeds and spreads.”276 Leopold only mentioned one problem exotic animal (a songbird) and it appeared, oddly placed, among a tersely worded table of predator-control references.277 A Year of Little Elton Books Across the pond, 1933 was a busy publishing year for Charles Elton. In addition to his summaries of the Matamek talks came two more small books. One of these was part of a multi-author series of monographs on biological subjects. The Ecology of Animals was intended as “ a brief but authoritative account of the present state of knowledge” for non-specialists.278 Something under half the length of his earlier Animal Ecology, it covered similar ground, but with a more practical emphasis. It was also the first of his books to allude, albeit in a tacit and indirect way, to mathematical theory, citing two works by Vito Volterra. Elton attributed Volterra’s importance to the association of ecology with economic problems, rather than with the mathematical methods Volterra had hoped “may be of some use to biologists.” 279 276 Ibid., 87-88 277 Ibid., 251 278 C.S. Elton, The Ecology of Animals (London: Methuen,1933) first edition dust jacket. 279 Ibid., 60. 125 The Ecology of Animals included several statements of interest regarding neobiota. The most notable among these had the opening elements of a ‘just so story’: “Originally the world had been split up by natural barriers into fairly welllimited zoogeographical areas. But the invention by man of better and better means of transport has had the unintended result of spreading round the world large numbers of animals whose arrival has often been the start of serious new pests or diseases.”280 This statement marked Elton’s first published mentions of the technogenic and unintentional aspects of such phenomena, both lending a hint of nostalgia for a less mechanized, more insular biogeography. Elsewhere in the book, Elton mentioned notorious introduced plants and animals of various sorts, including agricultural pests, as well as A.D. Middleton’s ongoing Oxford studies of American grey squirrels.281 Elton’s second 1933 book was titled Exploring the Animal World. Less technical than any of its precursors, it derived directly (nearly word-for-word) from a series of radio talks solicited of Elton by the BBC. As was typically the case when faced with the prospect of large audiences, Elton was initially unenthusiastic. But this time he soon relented. The last of four subjects he initially proposed was “introduced animals” (naming four) “and also possibly some plants” (naming one).282 He titled this segment, and its corresponding 280 Ibid., 77. 281 Ibid., 60, 72; see also A.D. Middleton, The Grey Squirrel (London: Sidgick & Jackson 1931). 282 C.S. Elton to M. Adams 31 January 1933. BBC Written Archives: Elton, Charles; RCont1, Talks File 1, 1932-1962 [BBC:ETF]. 126 chapter, “Plagues of Animals;” it was his first attempt to explain his views on the subject of neobiota to a general audience. In it he outlined several of the cases that later appeared in The Ecology of Invasions. His radio-enhanced reputation also supported an ‘op-ed’ article in the Times of London, titled “Alien Invaders,” which appeared almost two weeks before the “Plagues of Animals” talk aired. The article’s title played on the familiar theme of H.G. Wells’ War of the Worlds in an attempt to elicit public support for a nationwide animal population census.283 Only a few years later Elton had limited success in coaxing the BBC to consider the spread of American grey squirrels in Britain as “news”, and thereby to encourage public participation in a nationwide squirrel census.284 One BBC staffer’s hastily scrawled reaction: “What the hell? … There’s a fat lot of news in it.”285 One noteworthy chapter of Exploring the Animal World concerned “Nature Sanctuaries.” Elton explained the appeal of wildlife protection; but he also made plain the gulf he perceived between humans and animals, and the responsibilities incumbent on humans. There is an attractiveness to me in the idea that the thousands of wild animal species that live with us on these islands have a right to go on living there, although when they become a menace to other things that we value, then we have to ignore these rights … There is such a fascinating interest and great beauty in wild animals and birds and insects that they must be kept and cultivated as an amenity, as well as studied from a material and scientific point of view. 283 Charles S. Elton, “Alien Invaders” The Times, 6 May 1933, 13-14. 284 C.S. Elton to BBC Director of Talks 16 March 1937; J. Green to C.S. Elton 19 March 1937; J. Green to C.S. Elton 1 April 1937. BBC:ETF (cit. n. 282). 285 (Name illegible) undated note, found attached to Ibid (19 March 1937). 127 Don’t imagine that the question of wild-life sanctuaries can be left simply to Government departments, or scientific societies, or city councils. It can’t. After all, every garden and wood houses several hundred different kinds of wild animals, and your attitude towards the animal world round you will help determine the animal world that your descendants will live with; just as your attitude towards other people will help to mould the politics and social structure of the future.286 Elton’s views on the rights of wild animals and their value to humans would persist for the rest of his career, but his dismissal of “Government departments” was to undergo a substantial metamorphosis. Leopold Lauds Elton, and “Propaganda” Becomes an Issue Briefly reviewing Exploring the Animal World for a 1935 issue of the National Audubon Society’s magazine Birdlore, Leopold gushed, “Until I read Elton’s recent book, I for one believed it to be impossible to present the science of ecology to laymen. Now it is clear enough that it was impossible only because the right man had never tried it.”287 In addition to anointing Elton “the right man” to bring ecology to the masses, Leopold exclaimed to his own narrowly focused audience: “[Elton’s] pages are completely devoid of propaganda or preoccupation with any particular animal or issue. This somehow enhances their value as propaganda for that attitude toward Nature which he himself epitomizes in one sentence of characteristic simplicity. ‘We ought,’ he says, ‘to feel continually 286 Charles S. Elton, Exploring the Animal World (London: George Allen & Unwin, 1933), 81-82. 287 Aldo Leopold Book Review: “Exploring the Animal World” by C.S. Elton. Birdlore, 1935. IN Flader & Callicott, The River of the Mother of God (cit. n. 182). 128 pleased and excited that so many kinds of animals are able to live with us on the same island.’”288 Leopold evidently thought that propaganda of one sort was more laudable than propaganda of another sort. But it could not be said that “propaganda” was an unfamiliar phenomenon in 1930s America, or that it was viewed in neutral terms. Leopold doesn’t seem to have been saying, simply, “our propaganda ‘good,’ theirs ‘bad’.” Rather, he seemed to be invoking a version of what George N. Wallace would later call “the authority of the resource,” that nature has a way of telling us what is appropriate, if its message is appropriately interpreted and applied.289 Taking this idea as a cue allows us to query Exploring the Animal World for examples of statements that help us understand Leopold’s idea of propaganda. The chapter, “Plagues of Animals” deals with neobiota, and provides ample material. One conceit Elton employed there is both amusing and to the point: If all the animals in the world had a world conference to decide how to regulate their affairs, they would probably have three main resolutions on the conference table (some of the animals would have to sit on the table too, but there would not be room for all). They would pass a resolution condemning the practice among human beings of always labelling animals 288 289 Ibid. According to George Wallace ["Law Enforcement and the `Authority of the Resource,'" Legacy: Journal Of The National Association For Interpretation 1(1990):4-8]: “Wild nature can be said to have its own authority. Nature has her own rules, operates in certain ways, and has certain laws; there are consequences when we violate that authority.” This idea plays the stick to Leopold’s carrot from “The Land Ethic”: “A thing is right when … ” 129 as friends or foes of man. In the first place, you can’t make sweeping generalizations like that. [example omitted]. Then the animal conference would pass a second resolution against introducing animals from one part of the world into another, on the ground that it was most disturbing to their lives and caused endless problems. They would point out that the triumphs of engineering transport which human beings are so proud of only have the effect of carrying animals from places where they are at home, to countries where they cause nothing but trouble. [examples omitted] The conference would move that ships and aeroplanes be abolished, or failing that, that they should be properly quarantined and disinfected, and especially that animals destined for zoological gardens, or parks, or fur-farms, should be very carefully inspected before they were allowed in. The third thing the conference would do would be to appoint a commission to inquire into plagues, how to forecast them, and what to do about them when they were coming. Two human members would be appointed to the commission, one a naturalist and the other an eminent Victorian. The Victorian would be put there as being an authority on the powers of abnormal multiplication of the human species … [neoMalthusian digression] … The world conference of animals would probably break up in confusion because some of them had multiplied so much while the meeting was going on.290 George Orwell’s Animal Farm was still a World War away, and Lewis Carroll’s Alice adventures more than a World War behind. Elton doted on Edward Lear, and (as mentioned) Beatrix Potter.291 He could not resist including absurd imagery like animal delegates sitting on the table, but his fable was straightforwardly purposeful. It is perplexing that Leopold excluded an exercise such as this from his definition of propaganda. Propaganda was something Aldo Leopold encountered regularly, and not only in terms of international relations. The early twentieth-century American 290 291 Elton, Exploring the Animal World (cit. n. 286) 106-108. Charles S. Elton, “Small Adventures,” unpublished ms. CSEP A33. As a child, Elton’s first book was Potter’s Squirrel Nutkin, in which squirrels captured and delivered other small mammals to an owl in return for the right to safely gather nuts. 130 conversation about nature was often a heated one. Different organizations with incompatible goals (e.g., resource extraction, conservation, and preservation) employed unguarded rhetoric against each other.292 Several popular nature writers came under scrutiny for excessively anthropomorphizing their accounts of animal lives, and arguments over credibility were common.293 Often the peacemaker, Leopold seemed to be able to navigate these shoals as well as anyone, managing to stay on communicative terms with disparate interests. Perhaps that reflected skills he acquired from his years in multicultural New Mexico. Travels to Germany and the Fruits of Scientific Racism In late 1935 Leopold took “the only overseas trip he ever made,” not to visit Elton, but to visit Germany, under a foundation grant to “study the various ramifications of Game Management in relation to forestry.”294 His Deutschsprechend upbringing in Wisconsin doubtless helped make such an opportunity possible and gave it multiple attractions. Leopold’s review of Exploring the Animal World was published while he was in Europe, and therefore almost inevitably penned prior to embarking, or at the latest while catching up on reading during the requisite train and ship journeys. Fortunately, Elton’s book was (as usual) small and eminently portable. 292 See, e.g., Stephen Fox, John Muir and His Legacy: The American Conservation Movement (Boston: Little, Brown, 1981). 293 See: Ralph H. Lutts, The Nature Fakers: Wildlife, Science & Sentiment. (Golden, CO: Fulcrum, 1990.) 294 Meine, Aldo Leopold (cit. n. 163), 351. 131 Curt Meine depicted a Leopold increasingly dismayed by his unfolding experience of Nazi Germany. The Nuremberg laws prerequisite to legally dehumanizing German Jews were announced and adopted in the midst of Leopold’s visit.295 A supplementary decree providing the details of that dehumanization appeared the day before he set foot back in the United States.296 But German concentration camps for political prisoners had already been operating for over two years.297 According to Meine, Leopold “likely learned of the dread reality of the camps” while staying with the family of a Jewish academic in Silesia.298 If Elton’s animal conference did not seem overtly propagandistic to him before the trip, one wonders whether it struck Leopold as being so after his experiences in Germany, where an increasingly nationalistic re-imagining of even the landscape was underway.299 It eventually became clear that Madison Grant’s The Passing of the Great Race had been adopted as a fundamental tract of Nazi philosophy. “Grant even received fan mail from Adolf Hitler, who praised Grant's book, Passing of the Great Race, as his ‘bible,’ and the book was invoked as evidence for the defense 295 Congress of the National Socialist Workers' Party. “The Reich Citizenship Law of September 15, 1935.” www.mtsu.edu/~baustin/nurmlaw2.html, accessed 5 February 2006. 296 Meine, Aldo Leopold (cit. n. 163), 360. 297 Austin, 1996. www.mtsu.edu/~baustin/holokron.html, accessed 5 February 2006 298 299 Meine, Aldo Leopold (cit. n. 163), 358 Gert Gröning and Joachim Wolschke-Bulmahn, “Some Notes on the Mania for Native Plants in Germany.” Landscape Journal 11 (1992), 116-126. 300 in the postwar Nuremberg Trials. 132 Although Leopold applied arguments to wildlife introduction that were virtually identical to Grant’s regarding immigration, Leopold’s biographers have not specifically engaged with this issue, and it remains to be seen whether any useful evidence exists to support such an analysis. Meine called Leopold’s trip to Germany “perhaps the single most important paradigm-shifting episode of any American conservationist of the 20th century.”301 Leopold returned from his journey concerned about avoiding the negative aspects (ecological and political) of National Socialist conservation, and about the prospects for eventual war in Europe. But he had also found value in the German concepts of Dauerwald (“permanent woods”) and Naturschutz (“nature protection”) that he adapted to his own work. Leopold published several articles on the topics of German forestry and nature conservation during 1936, and reviewed a book by one of his German acquaintances in 1937.302 After that bout, Germany faded from his writings. 300 Dirk Olin and Edwin Black, “The Origins Of Hate” The American Lawyer 21 (2 September 2003), accessed online 27 September 2006 at www.waragainsttheweak.com/offSiteArchive/law.com/index.htm; The Avalon Project at Yale Law School, “Nuremberg Trial Proceedings Volume 18,” www.yale.edu/lawweb/avalon/imt/proc/07-10-46.htm accessed online 27 September 2006. 301 Curt Meine, “Aldo Leopold and the Huron Mountain Club” (anon. interviewer). Environmental Review 9 (2002),1-8. 302 Flader & Callicott, River (cit. n. 182) 360-362. 133 Parrying Xenophilia Introducing Leopold’s early 1938 essay “Chukaremia,” Flader and Callicott wrote, “As Leopold came more and more to appreciate the inner workings of the natural system, he had less and less patience with the imported species that were the pride of many state game departments.”303 This assertion appears problematic in light of Leopold’s explicit rejection of “mixing trout” two decades earlier. The chukar is an Asian partridge, and Leopold made it the namesake of an invented disease that “distorts the sportsman’s point of view,” leaving the impression that “foreign game birds are the answer to the ‘more game’ problem.” Focusing on birds introduced for hunting, Leopold listed twenty more species and invoked a further “score,” only two of which he considered even qualified successes. This very short piece took broad aim against the exoticism of sportsmen and game managers who failed to “face the question of doing something real for the game species already in our coverts.”304 A rare opportunity to actually prescribe management practices for a forest came Leopold’s way in late 1937, when the private Huron Mountain Club invited him to “devise a land program” for its 15,000 acre Michigan holdings.305 He divided his August 1938 “Summary of Recommendations” into six sets of statements regarding research needs and management emphases. The fifth set 303 Ibid. 304 Aldo Leopold, “Chukaremia” [1938], Ibid., 245-246 305 Meine, Aldo Leopold (cit. n. 163), 385. 134 evidences disdain for neobiota among a tersely-worded (and therefore slightly ambiguous) litany: “Exclude further roads; plant no more exotic species; tighten fire control.”306 A Honeymoon, A Shack, and a Doubtful Tale Married less than a year, Charles Elton and poet Joy Scovell traveled to North America in the early autumn of 1938.307 After a stop at the University of Wisconsin, they spent the weekend at the Leopolds’ “shack,” a cabin (née chicken house) on a worn out farm along the Wisconsin River near the town of Baraboo, about forty miles from Madison.308 Leopold acquired the land, in part, to test his ideas about ecological restoration, and his family spent many weekends there, documented by a handwritten journal.309 Leopold’s notations for this visit lit the participants as Aldo, his wife Estella, their daughter Estella (11), and sons Carl (18) and Luna (nearly 23), along with “Mr. and Mrs. Charles Elton.” Leopold recorded: “Highest water since 1881” occurred last week, according to neighbor Gilbert,” along with a few descriptive details. Other headings for the weekend included “Trees” (acorns planted), “Birds” (a short list of sightings), “Phenology” (seasonal botanical phenomena), and lastly, “Elton 306 Aldo Leopold, unpublished report. “Huron Mountain Club, Summary of Recommendations” ALP. 307 Charles S. Elton, diary of a visit to North America, 1938. CSEP C.24. 18-22. 308 Baraboo Wisconsin bills itself as the birthplace of the Ringling Brothers’ and other circuses, and as home of the World Circus Museum. Neither Baraboo, its Chamber of Commerce, nor Sauk County, nor the Sauk County Historical Society touts the Leopold connection; in fact, he is mentioned only in passing (if at all) on any of their websites. 309 Meine, Aldo Leopold (cit. n. 163), e.g., 364-365. 135 Tree Planted a seed shot of red oak and black oak acorns in front of shack for the Eltons.” 310 Elton also kept a journal of the trip. His report concentrated on aspects of the weekend that Leopold doubtless took for granted: “Rose early and after camp breakfast visited near-by oak and pine woods. This meant wading through remnants of an unusual summer flood and being badly bitten by mosquitoes of a vicious kind. Collected acorns of the red oak to plant in an Oxford College garden.”311 Leopold reported on the fortunes of the Elton oak(s) at the Shack in later correspondence with Charles. And as of this writing, there remains one Elton Oak on the property, the others having been culled to make room for what could become a very large tree. Contrary to expectations raised by the journal entries, the survivor is a (white) bur oak, Quercus macrocarpa.312 A passage in A Sand County Almanac makes it clear that bur oaks held a particular, and particularly Wisconsin-ish significance for Leopold. “He who owns a veteran bur oak owns more than a tree. He owns a historical library, and a reserved seat in the theater of evolution. To the discerning eye, his farm is labeled with the badge and 310 Aldo Leopold, “Shack” journal. ALP 9/25/10-7 003 001. 311 Elton, diary. (cit n. 307), 19. 312 Personal communication 2 February 2006, from Buddy Huffaker, Executive Director of the Aldo Leopold Foundation 313 symbol of the prairie war.” 136 Whatever they represented for Elton, the acorns he pocketed met a very different end, which is discussed in Chapter Four. A Concrete Creek, A Chinese Elm, Technophobia and Naturschutz Shortly after Elton’s visit, Leopold produced a lecture outline titled “Economics, Philosophy and Land.” He listed Elton among the “rebels” who “broke the laboratory ecologists’ unscholarly taboo on ‘practical’ problems.” Following several data slides, he listed among his “deductions”: “Many substitutions of exotics. Grains, grasses, weeds-- now trees and birds. The role of exotics in future systems is unpredictable.”314 A few months later, early in 1939, Leopold gave a public talk titled “The Farmer as a Conservationist” in which he idealized the properly managed farm. He punctuated his message with a closing jab at foreign landscape plants as a symptom of the unacceptable dominant paradigm: “It’s a wonder this farm came out without a concrete creek and a Chinese elm in the yard.”315 Later that year, Leopold spoke to a combined meeting of the Society of American Foresters and the Ecological Society of America [ESA].316 He more or less summarized his understanding of ecology in the process. A really thorough analysis of the talk lies beyond the scope of this paper, but some discussion is 313 Aldo Leopold, A Sand County Almanac, with Sketches from Round River. (New York: Oxford University Press, 1966). 30. 314 Aldo Leopold, unpublished outline, “Economics, Philosophy and Land.”ALP 9/25/10-6 016 006. 315 Aldo Leopold [1939] in Flader & Callicott, River (cit. n. 182) 255, 265. 316 Flader & Callicott, River (cit. n. 182), 266. 137 needed. Flader and Callicott asserted, “From the most recent contemporaneous ecological theory it abstracts an emerging new portrait of nature.”317 Exactly how it would have impressed members of each attending organization at first hearing is not clear. Opening with ecology, Leopold credited no other authorities, so he was not congratulating anyone present in the room. His choice of terminology, e.g., the “biotic organism” might not have satisfied Arthur Tansley, whose “ecosystem” had entered the conversation four years prior.318 Taking the pyramid of numbers as his cue and recasting it (interchangeably substituting “land”) as an energy circuit (per Alfred Lotka), he described the consequences of human technology to that circuit in woeful terms. “Man’s invention of tools has enabled him to make changes of unprecedented violence, rapidity and scope.”319 Violence against land by technologically adept humans themed the remainder of the piece. Human agency in creating neobiota was among the first counts in the indictment. “One change is in the composition of floras and faunas … Domesticated species are substituted for wild ones, and wild ones moved to new habitats. In this world-wide pooling of faunas and floras, some species get out of bounds as pests and diseases, others are extinguished. Such effects are seldom intended or foreseen; they represent unpredicted and often untraceable readjustments in the structure.” Further along: “The process of altering the 317 Ibid. 318 A. G. Tansley, “The Use and Abuse of Vegetational Concepts and Terms.” Ecology 16 (1935), 284-307. 319 Leopold [1939] in Flader & Callicott, River (cit. n. 182), 269. 138 pyramid for human occupation releases stored energy, and this often gives rise, during the pioneering period, to a deceptive exuberance of plant and animal life, both wild and tame. These releases of biotic capital tend to becloud or delay the penalties of violence.” Pausing to sum up, he revisited three points, the second of which was: “The native plants and animals kept the energy circuit open; others may or may not.” Reducing yet further, he asked, “These ideas, collectively, raise two basic issues: Can the land adjust itself to the new order? Can violence be reduced?” 320 These sentiments “saw and raised” Elton’s earlier contentions regarding technogenic mishaps. Leopold then made a set of (even then) hotly debatable and (at best) naïve claims.321 “In western Europe … the soil is still fertile, the waters flow normally, the new structure seems to function and persist. There is no visible stoppage of the circuit … Western Europe has a resistant biota. Its processes are tough, elastic, resistant to strain. No matter how violent the alterations, the pyramid, so far, has developed some new modus vivendi which preserves its habitability for man and for most of the other natives.”322 Three-quarters through, Leopold took a sudden turn with a one-sentence paragraph. “Forestry is a turmoil of naturalistic movements.”323 Now, three months after German troops had marched into Prague, and a month after Germany and Italy forged the “Pact of Steel,” he trotted out Dauerwald and 320 Ibid., 266-270. 321 See chapter three. 322 Leopold [1939] in Flader & Callicott, River (cit. n. 182), 270. 323 Ibid., 271. 139 Naturschutz, the latter in an interesting comparison: “The wilderness movement, the Ecological Society’s campaign for natural areas, the German Naturschutz, and the international committees for wildlife protection all seek to preserve samples of original biota as standards against which to measure the effects of violence.324 Intruders, Invaders, and a Romp Down the Rockies In 1941, with “Wilderness as a Land Laboratory,” Leopold coined the term “intruded animals” for wild horses living in a Mexican wilderness area, but it failed to penetrate the lexicon. About the same time came “Cheat Takes Over,” an article for Outdoor America that was later included in A Sand County Almanac. The “cheat” in question was downy chess or cheatgrass (Bromus tectorum) an annual grass native to the Mediterranean region. Leopold opened by describing a nameless fear that Daniel Simberloff would later label “invasional meltdown.”325 “Just as there is honor among thieves, so there is solidarity and co-operation among plant and animal pests. Where one pest is stopped by natural barriers, another arrives to breach the same wall by a new approach. In the end every region and every resource get their quota of uninvited ecological guests.” Since the standard version of the proverb states that there is no honor among thieves, Leopold’s intentions are either muddled by the misquote, or obscured if it was an intentional parody. 324 325 Ibid., 272. Daniel Simberloff and Betsy Von Holle. “Positive Interactions of Nonindigenous Species: Invasional Meltdown?” Biological Invasions 1 (1999) 1, 21-32. 140 Since the essay is readily available, an analysis suffices. There was a dual thesis: cheatgrass was aptly nicknamed, and its presence in North America was a symptom of management failure. In addition to the title character, nine relatively familiar examples of abutted biota were invoked: three each of plants, plant diseases and vertebrate animals. Concentrating mainly, but not exclusively on the grass, Leopold described their activities and effects using tropes that would then have rated blue-penciling if submitted to a serious journal. In addition to arriving uninvited, they (in alphabetical order) “blockaded” ducklings, “breached” walls, “choked” other plants, “found” seedbeds, were “inferior”, “landed via the back door”, “made a living on the roof”, and “romped down the Rockies.” Counts in the indictment included dense growth, inflammability, inedibility, invading, and prickliness. For benefit of the ladies, cheat was a predicament to wearers of “low shoes” or nylon stockings, and it “covered ruined complexions” of hillsides “with ecological face powder.” His browbeating final paragraph began, “I listened carefully for clues whether the [American] West has accepted cheat as a necessary evil, to be lived with until kingdom come, or whether it regards cheat as a challenge to rectify its past errors in land-use. I found the hopeless attitude almost universal.” Leopold acknowledged that Bromus tectorum had some positive attributes, but he made clear his judgment that these were lesser positives than the ones he attributed to native range plants.326 326 Aldo Leopold, A Sand County Almanac (cit. n. 313) 154-158. 141 Wartime Nationalism and the Proper Custodian Six months after the Japanese naval raid on Pearl Harbor, Leopold raised a Dauerwald-based alarm of his own against the reckless wartime harvest of oldgrowth hardwood forest in Michigan, reflecting his own wartime nationalism: “The sugar maple is as American as the rail fence or the Kentucky rifle … we make shift with substitutes. The poorest is the European ‘Norway maple’, a colorless fast-growing tree persistently used by misguided suburbanites to kill lawns. Wisconsin has used Norway maples to shade its capitol. No governor and no citizen has protested this affront to the peace and dignity of the state.”327 Treading more carefully, he made an example of a forest “covering a mountain on the north flank of the Alps” without ever quite mentioning that the mountain was in Germany.328 Later the same year, Leopold mentioned neobiota among a list of “overemphasized” activities and facilities: “game farms, fish hatcheries, nurseries and artificial reforestation, importation of exotic species, predator control and rodent control.” Nevertheless, he pinned his hopes on the “real and indispensable functions of government in conservation. Government is the tester of fact vs. fiction, the umpire of bogus vs. genuine, the sponsor of research, the guardian of technical standards, and, I hasten to add, the proper custodian of land which … is not suited to private husbandry.”329 327 Leopold [1942] in Flader & Callicott, River (cit. n. 182), 292. 328 Ibid. 329 Ibid. 300. 142 Most of Us Have Been on Rats Just then, Charles Elton’s Oxford Bureau of Animal Population was busily emphasizing rodent control. In a November 1942 letter, Elton reminded Leopold of this fact and gave him a progress report: You may remember my telling you that when Prague was raped, I decided to get busy on plans for war research, as it was plain that the Govt. had decided to reserve all of us University scientists anyway, but had no particular notions what to do with us then! So we got a flying start, in September 1939, and have grown into a team of about a dozen men, most of whom have made themselves sufficiently useful to the Govt. to be retained on the job. My first thought was to get [Doug] Middleton back, and he has turned up trumps, as one would expect … In the first year of the war he had covered and solved the problem of rabbit control on farm-land (not woodland and scrub, tho’), and by the third year of the war the rabbit has become a rare country species over much of the better farmland of England … Now he is on the rural brown rat, and is perfecting a method of getting 100% kills with poison baits. Most of us have been on rats, one way or another, and [Mick] Southern is on house-mouse baiting, which is a serious item in our long-term stores … More rats are being killed than ever for years! This winter, it’s a drive on cornstack rat poisoning, which we only got the answer to last year.330 Elton’s letter is a window into his wartime world, which was evidently a busy one. He opened it with an apology. “Dear Leopold, I am pretty hopeless as a correspondent, and yet the pile of your letters and reprints [including “Cheat Takes Over”] have lain on my desk ever since I last wrote to you. I have yours dated 14 Feb. and 25 Nov. 1941, for which we were both most grateful.” A scientist with rationing on always on his mind, Elton eschewed the idea of unnecessarily duplicating well-established results: “At one point I thought of writing you a thorough account of what life is like here in wartime, but I expect this has been so relayed by U.S. men over here, as to be hardly worth while 330 C.S. Elton to A. Leopold 18 November 1942. ALP 9/25/10-8 001 002. 143 now.” He was hardly immune to war fever: “It is a grand sight to go up to London and see the amazing mixture of men and women from all countries of the Empire and U.S.A. and various small European units too. One does get the feeling that here have gathered a number of essentially angry people from every part of the world, to take part in a thorough clean-up.” Evidencing his dissatisfaction with the conclusion of the previous conflict, he continued, “And this time they won’t stand for any half-measures, either in the military sense (or what will be just as important) in the social scene after the war.” Elton even betrayed some personal exoticism, if perhaps in terms too stereotypical for the comfort of later readers: “It is quite a portent to see convoys of negro troops from your country, drive smiling through the quiet English country villages!”331 (The same year, in his book Voles, Mice and Lemmings, he routinely differentiated between arctic foxes and red foxes using the aptly descriptive but imperially burdened terms “white” and “coloured.”)332 In closing, Charles assured Aldo, “We very often talk about the Leopolds and our other American friends, and wonder whether Catherine [born 1940] will be big enough to travel out when the time comes for us to revisit the U.S.A. We would be very interested to know how your family is faring, and whether the war has swept any of them into the service yet.”333 Carl Leopold had enlisted in the 331 Ibid. 332 Elton, Voles (cit. n. 247), 258-320. 333 C.S. Elton to A. Leopold 18 November 1942. (n. 330.) 334 Marines on 8 December 1941. 335 Luna was in the Army. 144 It had been a year since Leopold’s last letter, and at least two since Elton’s. Conservation in Great Britain: Only the State With his 1942 letter, Elton enclosed a copy of “a memorandum I wrote the other day. It was for A.G. Tansley, who is running a committee of the British Ecological Society on the general question of nature reserves.” Reflecting both the modern disciplinary divide and his penchant for sharp remarks, he quipped, “It is intended to make a group, largely of botanists, faintly aware of the change that has come over conservation ideas.”336 Titled “Conservation of Wild Life in Britain,” its seven-plus single-spaced typed pages laid out Elton’s justifications and vision for a government-protected, but likewise government-controlled ecology. In itself, the document is fodder for an extended discussion. It seems to show Elton unburdening himself of long-incubated but previously unarticulated ideas. The analysis here necessarily concentrates on issues of neobiota, and on identifying conservation ideas consistent with Leopold’s. But it made plain that Elton, while working under government aegis, had carved out a major niche in his conservation philosophy for “government departments.” The document gives every impression of representing a barely revised stream of consciousness. It was divided into ten major, numbered paragraphs; numbers six, seven, eight and ten had demarcated subsections. Subparts of 334 Meine, Aldo Leopold (cit. n. 163), 420. 335 Ibid., 432. 336 C.S. Elton to A. Leopold, 18 November 1942. ALP 9/25/10-8 001 002. 145 paragraphs six, eight and ten were headed with lower-case letters, but paragraph seven’s with Roman numerals. Paragraph nine was, at some point, designated “9½” but the numerals (only) of the “½” character were individually struck out; it was also titled “Appendix 1. The U.S. Fish and Wildlife Service” and included a list of enclosures, most (but not all) produced by that agency. Paragraph ten was titled “Appendix 2. Notes on British Mammals.” Only nine of its thirteen discrete topics were subtitled. Various typographic errors were corrected by hand, as if it were a penultimate draft. Elton also wrote in terse marginal comments, some obviously for Leopold. Elton gave no indication that it arrived on Tansley’s desk in the same form, but also made no suggestion otherwise. Paragraph one laid out two assumptions, “that conservation of animal life is desired” and that “controversy about cruelty to animals is excluded from consideration.” Paragraph two justified “measures for the preservation of the native fauna.” It centrally cited “the increased chemical and mechanical destructive power of people (either for destroying the existing environment, especially the vegetation, for various purposes, or by destroying the animals with efficient methods [listed] evolved in the last 150 years.” Paragraph three explained that “the motives for such preservation are very mixed, often indeed flatly opposed to one another,” and “It is unusual for any single person to like all animals.” It gave three examples of selective or conflicting valuation.337 337 C.S. Elton, unpublished memorandum, “Conservation of Wild Life in Britain,” [und., 1942] ALP 9/25/10-8 001 002. 146 Paragraph four opened “Much of the controversy that arises about preservation, is not over the question ‘Is it a good thing to preserve animals’, but over which species shall be given priority in the scheme of things that each particular individual favours. That is why conservation is so essentially an ecological expression of the resultant of many warring philosophies. The ecologist, or the scientist generally, can hardly expect to impose all his views about which species are desirable, and in what proportions; but he can point out several principles that will lead to a more rational and successful solution of the problem, and he can often, already, point out that the results expected even by the proponents of particular views are different from those that will really occur.” The remainder of paragraph four provided a similarly difficult-to-follow example using rabbits. 338 Paragraph five was a sort of conclusion to the previous four, but not an immediately self-evident one. Here, in full: “In this memorandum I shall therefore chiefly deal with the sequence that animal life cannot be conserved simply by a passive policy of fencing off certain areas of land as reserves, that the problem always eventually becomes one of attaining a certain density and distribution of population for the species concerned, and that this means active and positive management, and management cannot be done without research information and experimental applications of this to the field problem.”339 338 Ibid. 339 Ibid. 147 Paragraph six had nine subparts providing a difficult-to-characterize array of “examples of the kind of mixed motives operating in the field of animal preservation.” Subpart (a) dealt with value conflicts surrounding small game, predatory and domestic animals, some of which were labeled “introduced.” Subpart (b) was a short statement (here in full): “A shooting man often protects badgers, although they are supposed to be enemies of game-birds (on little evidence). He happens to feel that the badger should be preserved.”340 Subpart (c) was confusing. In full: “The red [native] squirrel is very widely looked upon as a charming animal, by people who take an ordinary interest in country or suburban scenes. He is one of our amenities. The grey squirrel, introduced on a ducal estate for amenity, may have a large responsibility for the widespread decrease of the red. The red squirrel has done shattering damage to Highland pine [introduced] forests by ring-barking below the crown, and is certainly a less serious forestry pest than the grey squirrel, at any rate in maturing conifer woods.” Subpart (d) explained a wartime shift in attitudes regarding rabbits due to the need to maximize production of foodstuffs; (e) concerned the single, discrete case of a biologist trespassing on private land. Subpart (f) provided two more examples of thoughtless zoology fieldwork. The second: “[a] biologist, on [a] Hebridean island, although he has a high consciousness of conservation, unintentionally kills most of the island mice, in 340 Ibid. 148 trapping [for museum specimens].” Handwritten here in the margin, with an indicating arrow: “This was me!”341 Paragraph 6(g) was wholly concerned with neobiota: “Different landowners have introduced, for the sake of amenity or sport on their estates, [seven mammal species, four birds, a reptile, an amphibian, and three fishes] to mention only some of the vertebrates. Fur-farmers have introduced [three smallish mammals] which have escaped, the former seriously; also [two predatory mammals]. In these cases the interest in the bizarre and new was stronger than that in the native species though the introducers probably thought that their actions would have no special effect on the native fauna, or flora. The man who introduced [one species] (which became a violent local pest by their noise, but subsequently failed to maintain their first breeding vigour) stated that his reasons were (1) their fine colours (he was an artist) (2) to prevent them being kept in a refrigerator for experiments.” Subparts (h) and (j) respectively concerned the attitudes of Cornish fishermen toward seals, and the ecological roles of voles. Elton omitted (i) from the sequence.342 Paragraph seven opened as a literal pivot. First it summarized: “Such examples show the tangled thicket of motives and habits and activities that affect the population size and distribution and prospects of wild vertebrates in Britain;” then, it introduced twelve “general principles” for “laying down policies of preservation” that “concern the mechanics of conservation, rather than the rights 341 Ibid. 342 Ibid. 149 and wrongs of conflicting ideologies.” With the exception of the first, each of the twelve subparts of paragraph seven opened with underlined phrases. Subpart I contrasted the strategies of preservation and conservation, likening the former to keeping a monument or a library, and the latter (which he favored) incorporating preservation with “promotion of interest by education and publicity, the balancing of material and cultural claims, research, and management.” Subpart II began, “Active forces of destruction require active measures of conservation,” followed by an assertion that by itself, designating preserves was an inadequate strategy. Subpart III began “The usefulness of a reserve for any particular species, is correlated with the size and range of movements of the animal, and with the vulnerability of its breeding population.” Its remainder consisted of examples of wide-ranging, colony-breeding and narrowly endemic species. Subpart IV: “In this country every section of population of a species is someone’s property i.e. the vast majority of animals are and will continue to be at the mercy of private people or the agents of corporations etc.” He expanded, but did not contrast the situation with public ownership of wildlife as codified in the United States.343 Subpart V employed III and IV as its premises. “The range of movement of British vertebrates is usually larger than the area controlled by one [human] occupier. Therefore the same animal may constantly pass through or live successively in, different bits of property,” to its recurring peril. Elton 343 Michael J. Bean and Melanie J. Rowland, The Evolution of National Wildlife Law. 3rd. Westport, CT: Praeger. 1997. 150 ambiguously invoked neobiota, and (inadvertently?) gave Leopold a poke regarding cats: “The shooting of valuable house-cats by [game]keepers, of herons by fishermen, the sore problem of pest reservoirs [with a list] and introduced animals are examples.” Then Subpart VI took a leap: “This situation of mosaic influences on animal life requires central State control, acting through regional units. Animal populations cannot, any more than the sick, the unemployed, or the aged, be left to an unguided struggle for existence. Only the State can focus a sufficiently representative body of opinions and scientific facts onto these complex questions, and arrange a distribution of animal life that will provide the maximum chances of survival for animals and of amenity for people, with the economic and epidemiological interests of the community. The present situation is one of complete ecological anarchy, with thousands of people doing one thing, and thousands doing another, often on the basis of completely wrongly based views of the nature of the problem. Two handwritten (apparently by Elton) marginal notations accompanied his call for a state-planned, welfare ecology. The first, perhaps, are the two letters “AL”; the second, “provision for L.” If meant to invoke Leopold, his intention remains obscure. As if recoiling from the political audacity of all that preceded it, subpart VII warned of scientific uncertainties. “Most wild populations are subject to changes in their populations brought about by natural causes. These are often attributed to human influence, but are afterwards found to be independent and largely mysterious changes in numbers [with examples]. Research has to be initiated in order to distinguish between human and natural causes of changing populations.” Subpart VIII wove together previous themes: “It is no good having a conservation policy that cannot be enforced.” Elton followed with reminders that 151 gaining public support for a program via “educational and publicity work of an impressive and penetrating character,” promising “in this way the public becomes its own wildlife guardians, with thousands of alert and informed representatives spread over the country.” Subpart IX completed the thought begun in VII: “It is no good planning a conservation policy without a solid basis of both ecological and social facts. Our work at Oxford on rodent control (which is a sort of mirrorimage of conservation) has taught us the intractability of man, as well as of the rat.” Elton pointed out the zoologists were too-often focusing on the wildest of the wild remnants, rather than the countryside at large. Then he brought the point home to Tansley: “Just as, if I may be pardoned!!, your wonderful book on British vegetation hardly mentions hedges, which form one of the chief habitats for animals in this country.” Subpart X appears intended to mollify his memo’s recipient after the criticism, by closing ranks with him and peering into their future. “The work of scientific research and advising is the function of ecologists. The ordinary zoologist in this country is still strikingly unconscious of dynamic ecology. The museum man’s day is rather over, in so far as the general taxonomy of British vertebrates is pretty well worked out.” Paragraph seven, Subpart XI was a longish discussion of organizational relationships that need not be detailed here; “Some State agency should be set up with the function of coordinating wildlife conservation, control and management in Britain. Of minor interest is the assertion, “This plan only follows the course recently adopted in the U.S.A.” Subpart XII is a short statement. “This organization would administer the wild life aspects of National Park and other 152 public nature reserves, as an essential part of a wider policy of the promotion of conservation.” Paragraph eight was a list of “Urgent conservation problems,” mostly related to specific animal populations on named small islands. The last two statements broke that pattern. He revisited neobiota and invoked scientific authority: “Forbid absolutely the introduction of foreign animals for any purpose, without the permission of the Wildlife Board [proposed in paragraph 7.XI], which would have overriding powers of prohibition. (This has now been done in Hawaii, where the Board consists mainly of scientists).” Coming last was a suggestion that, should a National Park be established, “animals should be allowed to migrate from protected source populations inside it to surrounding areas.” Paragraphs nine and ten (the “appendices”) respectively described conservation-related agencies in the United States (with one significant error, placing the Forest Service in the Interior Department), and miscellaneous notes reminiscent of the list in paragraph eight. Several of the latter mentioned neobiota, most specifically, subpart “(g) Introductions.” Three ungulates were “established from parks and spreading. [One of these] was, before the present War, becoming abundant on the Dorset and Hampshire heathy lands.” Turning to what seemed his perennially favorite case, “Muskrats introduced in many furfarms, and escaped in Shropshire, the South of England and Scotland, have been wiped out at a cost of well over £20,000. In the course of Scottish trapping enormous numbers of water voles and some wild birds such as coot and moorhens were accidentally destroyed.” Another rodent “continues to escape at 153 intervals, but is fairly easily killed. It is, however, wild in France and parts of southern U.S.S.R.” and a third “introduced … some fifty years ago, is apparently widespread, but in a cryptic way owing to its elusive habits and superficial resemblance to [a better-known introduced rodent]. It is a potential fruit and nut pest.” Elton revisited another favorite case for the second time in the same memo: “The American grey squirrel continues to spread, and probably to oust the red squirrel,” and he finished with “Black rats are spreading inland from coastal towns, but it is hoped that control of this and the brown rat [referring to the BAP’s then-ongoing program] are well forward and will be successful in the next ten years.” Finally, after discussing bat management problems, Elton terminated his memo with unexpected imagery that rivaled his best efforts elsewhere. “The sight of a stream with Daubenton’s bats weaving silently over it at dusk, is one that anyone should wish to see.”344 However peculiar it was, Elton’s memorandum to Tansley gathered aesthetic, ethical, scientific and economic justifications for wildlife conservation without really sorting or reducing them into those categories per se. Thus it restated the ideas Elton had expressed a decade earlier through the mouths of his “world conference of animals” and elsewhere in Exploring the Animal World. His had begun emphasizing the role of government, but still stressed the importance of cultivating public goodwill. Some, but certainly not all, sections of this memo were incorporated with little editing into the May 1944 British 344 Elton, “Conservation.” (cit. n. 337). 154 Ecological Society committee report “Nature Conservation and Nature Reserves.” 345 A Paradise, Lost? Most likely within a month of firing off his memo and his letter to Leopold, Elton submitted a polemical essay to an obscure, refugee-run pseudo-journal, Polish Science and Learning.346 Titled “The Changing Realms of Animal Life”, it was published in February 1943. Three of its four pages explained Elton’s conception of prehistoric biogeography and the fourth was pointedly alarmist. He reiterated a favorite assertion from his Oxford lectures on animal distribution: “we are … present at the first stages of a historic and irreversible breaking down of the great realms of plant and animal life, evolved over a hundred million years or more. This change has profound implications for the material and cultural environment of our activities, for many hundreds and even thousands of years to come.” Elton imputed a suite of motives for intentional animal introductions including: “a desire for acclimatization of familiar native forms in a strange land”; “the desire for sport or food or fur”; that “the familiar species…are not enough satisfaction to [someone’s] need for the bizarre”; and “the need to control some pest already accidentally introduced.” He revisited his decade-prior call in the Times, for coordinated science and government action, and echoed and amplified his Tansley memo: “A hundred years of modern transport has brought 345 British Ecological Society. “Nature Conservation and Nature Reserves.” The Journal of Ecology 32 (1944):45-82 346 An editorial in Polish Science and Learning 2 (1943) 5 begins: “The Ministry of Supply has refused a licence to publish [PS&L] as a periodical. We shall therefore keep our publication to a SERIES OF BOOKLETS.” 155 about an ecological pandemonium which nothing but the most thorough and careful research and international co-operation can handle successfully.” 347 It is not far-fetched to suppose that Elton, steeped in literary allusion, knew perfectly well that “Pandemonium” was the name Milton coined for the capital of Hell in Paradise Lost.348 Elton promoted “The Changing Realms of Animal Life” with superfluous comments in a 1944 book review for The Journal of Animal Ecology. There, too, he closed with militaristic, even cataclysmic warnings: These are major engagements in a violent struggle against the spread of undesirable plants and animals that is affecting every country. As I have pointed out elsewhere [Elton 1943] we are witnessing not only the immediate dislocations caused by the introduction of various species into countries new to them, but a vast historical event — a zoological catastrophe, which is the beginning of the breaking down of Wallace’s zoogeographic realms and innumerable island isolations by the activities of man, a process which will eventually reduce the rich continental faunas to a zoned world fauna consisting of the toughest species … This constant bombardment with foreign species is illustrated [in the item being reviewed] … It is clear enough that the price of ecological stability (if one can dignify our fluctuating natural communities by this term) is eternal vigilance and a high sensitization of administrators everywhere to the ecological results of efficiency in modern transport.”349 347 Charles S. Elton, “The Changing Realms of Animal Life,” Polish Science and Learning 2(1943)7-10. 348 John Milton, Paradise Lost (1667) Chapter 10: “about the walls Of Pandemonium; city and proud seat Of Lucifer.” 349 Charles S. Elton, “The Biological Cost of Modern Transport,” The Journal of Animal Ecology 13 (1944), 87-88. Since Elton edited the journal in which this review appeared, there was nothing to check such outbursts other than his own recognizance. 156 What is a Weed? and Pheasants Under Scrutiny Dated 2 August 1943 but unpublished until 1991, “What is a Weed?” was Aldo Leopold’s very belated (when penned) review of a 1926 Iowa Geological Survey bulletin, The Weed Flora of Iowa.350 It stands as anomalous among his writings, in that three of the eleven species he singled out as unfit for persecution at government behest were also examples of neobiota. He even closed ambiguously. “It seems to me that both agriculture and conservation are in the process of inner conflict. Each has an ecological school of land-use, and what I may call an ‘iron-heel’ school. If it be a fact that the former is truer, then both have a common problem of constructing an ecological land-practice. Thus, and not otherwise, will one cease to contradict the other. Thus, and not other, will either prosper in the long run.”351 Dated 7 April 1944, “The Ringnecked Parvenu” appears to be an unpublished, three paragraph essay fragment.352 It exists in two versions among Leopold’s archived papers, a handwritten manuscript with additions and deletions, and a derivative typescript.353 I reproduce it here in full, including insertions (underlined) and struck-out phrases: 350 Weed Flora’s lead author was L.H. Pammel, perhaps notable for being the first (1893) to author a book in English with ecology in the title (McIntosh, Background, 29). 351 Aldo Leopold, “The Ringnecked Parvenu,” unpublished ms. (1944) ALP 9/25/10-6 016 007. 352 A “parvenu,” by definition, is a nouveau-riche upstart, lacking the refinements of the social class to which he (or she) pretends. 353 Respectively, ALP 9/25/10-6 016 007 and 9/25/10-6 017 008. such typescripts were prepared by Patricia Murrish, secretary of the University of Wisconsin 157 The pheasant, among ecologists, is held in low esteem. He is a bird, yet something less than a bird; he is successful, but in places too much so. He is suspected, perhaps guilty, of displacing his native counterparts: the quail, the prairie chicken, and the ruffed grouse. To enthuse about pheasants often ellicits [sic] that silent rejoinder, the lifted eyebrow, and implies more interest in gunpowder than in conservation. I share all these prejudices, against pheasants and pheasant enthusiasts, and yet at times I wonder whether they do not arise from the fact that the pheasant is too much like ourselves. After all he only did what we did: he appropriated a new land by outcrowing, outeating, and outbreeding its native inhabitants. He, and we, are parvenus. He, and we, are mongrels. He, and we, are brash, disputatious, noisy, too enamored of the world’s goods, and too successful in acquiring them. The pheasant has virtues which we perhaps lack. The mallard, the wild turkey, and the domestic fowls, in bowing to the necessity of living behind chickenwire, have lost their capacity to live otherwise. He But the pheasant is tenacious of his wildness; he alone, of all the game birds, endures generations of soft living without losing his capacity to survive in the wild, or his preference for doing so. It is needless to elaborate on the points Leopold made, except to clarify that the epithet “mongrel” (as applied to pheasants) reflects the American introduction of several recognized subspecies of Phasianus colchicus from populations in various regions of Eurasia, and of deliberately induced crossbreeding among them before release.354 And it is worth noting that the Department of Wildlife Management, when the papers were organized for preservation in 1948. (University of Wisconsin, Madison. Archives and Records Management: Aldo Leopold Papers finding aid. http://digital.library.wisc.edu/ 1711.dl/WIArchives.LeopoldPapers), In this case, the typescript reflected the handwritten corrections; but the still-legible deletions on the manuscript significantly affect the reading of the piece, and demonstrate the author’s attitude and intention at the time of writing. Since Leopold’s handwriting was quite legible, and the original was also preserved, the purpose of preparing the typescript is unclear. 354 Christopher Lever, Introduced Birds of the World, (Harlow, UK: Longman Scientific & Technical, 1987),150-175. 158 concerns Leopold expressed in 1944 echo strongly among the doctrines of invasion biology some sixty years later.355 I Started this Letter Nine Months Ago; Of Quiet Visits Evidently responding to some unpreserved but discouraged words from Aldo, Charles began a cheery typewritten rejoinder on 24 January 1945: “Dear Leopold, It is always delightful to get any letter from you, and the reprints from you and your Dept., and you must never think that my poor reply is a measure of my interest in Leopold Activities.” The balance of the first page was all bragging on rat control. Page two faltered. “So far so good. But this year we all feel rather flat and tired, and I am facing the problem of rehabilitating these people for postwar longer-term research. On the whole the ‘reserved’ scientist works over-hard to justify his reservation from military service. Last autumn I lost [laboratory assistant, Richard] Ranson, at the age of 33, because his heart cracked up. He was a brilliant man, in spite of having no higher training beyond a long apprenticeship with research men. I miss him very much: he was to me as much as [Paul] Errington or [Errington’s students, Frederick and Frances, the] Hamerstroms would be to you.” Ranson had first worked with Elton in the late 1920s, and was among the first staff members of the BAP. Crowcroft called his 355 See, e.g., Kay Milton, Loving Nature: Towards an Ecology of Emotion (London: Routledge, 2002); Mark Sagoff,. What's Wrong with Exotic Species? In Philosophical Dimensions of Public Policy, edited by V. V. Gehring and W. A. Galston. (New Brunswick: Transaction 2002); Subramaniam, "The Aliens Have Landed!" (cit. n.10); M.K. Chew and A.L. Hamilton, “Nativeness Considered,” in prep. 356 loss to the group “grievous.” 159 Coincidentally, Richard Ranson and Geoffrey Elton had died at very nearly the same age. There the typing ended. Elton continued by hand, entering a new date, and then: “It’s no use concealing the fact that I started this letter nine months ago. The world seems to have whizzed round at a remarkable pace since, but we are still here and flourishing.” World War Two had ended in both theaters between the inception and completion of one letter, which now wound around many topics, as if making up for lost time. Elton proffered family statistics (“Catherine is 5½ … Robert 1¾”) and sweeping political generalizations (“What we really need in this country are more houses, less work for the housewife, and a smaller national debt.) He hinted at new support for his hypothesis that competitive exclusion was most evident at the subgeneric level. Rhetorically, he asked, “Don’t you think we must all resist the deluge of ad hoc work & just sit back and think? There are too many busy and well-intentioned vampires wanting to sip our good blood.” 357 By that time Leopold had been working toward the book of collected works that would become A Sand County Almanac, sending both new and revised essays to trusted friends, including Elton. Here he reacted: “You have sent me such good letters, notes and papers, that I can scarcely begin to catch up in answer & comment. Life has been thick and hard and spare time very scarce. This summer my old father died at 84, after a brief illness, which was a sorrow to 356 357 Crowcroft, Elton’s Ecologists (cit. n. 184), 6; 44. C.S. Elton to A. Leopold 24 January/10 September 1945. ALP 9/25/10-3 001 002 160 us & also landed me in a spate of business.” After detouring into a request for help acquiring a reprint from one of Leopold’s colleagues, Elton returned to the previous thought and concluded: “Let me know how your new book is going on. Oh for a non-pedestrian work on the subject! I shall look forward eagerly to yours.”358 In 1945, three books bearing on conservation planning for postwar Britain were published. Elton reviewed them all for the November issue of The Journal of Animal Ecology. One was A.G. Tansley’s Our Heritage of Wild Nature, which built directly on the conservation planning efforts of the Wild Life Special Committee, and Elton’s 1942 memo. Incidental commentary was becoming a regular feature of Elton’s reviews, and this time he began by quoting selections from Thoreau in support of his contention that “Wildlife is not only a pleasure, a resource or an intellectual problem. Given a reasonable chance, it simply exists and abounds in its own extraordinary variety and rhythm, whether we write about it, eat it, try to measure its pulse — or not.” He also made a curious, historically naive observation: “But [the quoted Thoreauvian passage] was written ninety years ago, before we had seriously begun to scramble together the floras, faunas and diseases of the continents [and develop various technologies].” In Elton’s view, neobiota had been insignificant before 1855, and perhaps until much later. He went on: Man is now a pervading ecological force, and the only way he can conserve even modified natural plant and animal communities is by taking direct responsibility for their survival, scientifically investigating their environment and 358 Ibid. 161 populations, and framing policies of management, if not for every species (of course an impossibility), at any rate for the habitats and most influential forms (including himself). Prof. [sic] Tansley … reviews the whole problem of conserving our native (and established alien) vegetation and animal life, which he sensibly includes all together under the general term ‘wild life,’ much of which he recognizes, however, to be only semi-natural in its community balance [emphases added].359 Without ever acknowledging the extent to which the book reflected his own contributions, Elton also wrote, “I do not know any book on ecology and conservation that has quite the qualities of this one: authoritative on the dynamic ecology of plant communities and well-informed about the British rural scene, equally aware of the special problems of animal populations, stating with massive wisdom the different values of material development, science, education, humaneness and enjoyment of rural beauty and bringing the threads together into a lucid exposition of practical policy for Government action.”360 Apparently Elton was satisfied by 1945 that he had made the eminent plant ecologist at least faintly aware of the changes that had come over conservation ideas. Elton wrote another lengthy and rambling letter to Leopold on 18 March 1946. After opening with an appreciation of Joy’s poetry, he turned to an appreciation of Leopold’s essays. “It was fine to get all the news in your letter of October 2nd [not found]. We both liked very much the descriptions in “Green 359 C.S. Elton, “Conservation Plans for Britain” The Journal of Animal Ecology 14 (1945) 156-158. 360 Ibid. 162 Lagoons” [a 1920’s canoeing adventure in the Colorado River Delta] which gave one both the vicarious sensuous pleasure of enjoyment of nature, and at the same time the almost intolerable sense of loss (that is at the back of my mind all the time) caused by Human Progress.” After some general thoughts on ways to affect government policies in Britain, he detailed the progress on that front. “The Wild Life Conservation Committee I am on is working to its Report this summer. I think we shall have then done about 40 meetings. The ecologists have rather assumed direction! (…) This is all really confidential still; but I don’t mind telling you we are gunning for a whole new Govt. unit with a highly trained scientific staff & run by scientists plus others.” Returning to a perennial complaint, Elton wrote, “I hope you will get some time to do more writing. The present time does not encourage contemplation and quiet industry. Everyone seems to want to consult everyone else!”361 After remarking on his satisfaction that Leopold’s boys had come safely through the war, Elton zigged to ecology, with a brief preview of a paper in press [ref]. He zagged to Joy’s poetry. Then he solicited Leopold’s views on American big game census methods, opined on the pointlessness of teaching the English taxa to ecologists destined for research in far flung corners of the empire, assented with Leopold’s views on “1080,” a poison then newly in use for rodent control, and a requested a replacement copy of a lost document. By page seven, he finally made what might have been his real point. “I don’t seem like getting abroad this year, feeling rather like pottering at home, and not at all like the 361 C.S. Elton to A. Leopold 18 March 1946 ALP 9/25/10-3 001 002. 163 impresario or even a modest research tour. I had a chance to go to the Swiss National Park, but have also sidestepped that. Next time I (or we) go to the United States, I would rather spend some quiet visits with only two or three friends, not do a high-speed tour. I hope the Elton oaks will not have grown too large before this happens.”362 A Large Number of Matters to Attend to The year 1947 opened with a rapid exchange of letters. On 2 January Leopold wrote to Elton regarding the “Committee on Foreign Membership, of the Wildlife Society.”363 Receiving it almost three weeks later, Elton replied “at once,” complaining about the state of the mails, and about his lumbago, but also providing a list of prospective foreign members in the U.S.S.R., Norway, Denmark, and Holland. He continued, “New Zealand is getting more active now that the Pole Wodjiski (or some such spelling) is at work. He … became Polish consul in N.Z. during the war & then was outed by the phony Polish Govt. there is now. An excellent & cultivated man, whom I know.” In the same context, Elton also joked, “Shall we be able to award ourselves travel fellowships to visit each other, or are our countries still allies and not ‘foreigners’??”364 Between mailing his 2 January letter and receiving Elton’s reply, Leopold was elected President of the Ecological Society of America — even though he 362 Ibid. 363 A. Leopold to C.S. Elton, 2 January 1947 ALP 9/25/10-3 001 002. 364 C.S. Elton to A. Leopold 20 January 1947 ALP 9/25/10-3 001 002; the name’s preferred transliteration was “Wodzicki.” 365 was not an active member. 164 Then, evidently feeling pressure from senior members to coax the reluctant Englishman across the Atlantic for an American appearance, he mailed two letters to Elton on 30 January. The first was oddly addressed to “Dr. Charles Elton.” Odd because neither Elton nor Leopold ever earned a Ph.D. and they shared a lifelong avocation of correcting those who initially addressed them as “Doctor.” Following a brief mention of old business, he entered a one-sentence paragraph: “At this writing, I wish to consult with you about an American trip.” The next two short paragraphs contemplated a symposium at a joint meeting of the Ecological and Zoological Societies the following December, “and also the financing of a trip for you.” Apparently, though, Elton’s dislike of such meetings was notorious. “They wish first, however, to have some assurance that you would consider such a trip favorably, if the where withal were obtained.” Then the tone changed, becoming gradually more informal. “Speaking now personally, I cannot urge a speaking trip on you in midwinter, because Chitty and Lack [see below] reported you as overworked and in need of a rest … [Two pending letters] are probably the forerunners of a whole swarm of invitations, once word gets around that you are coming. In short, there is a conflict in my mind between personal consideration for you, and my ‘official duty’ to the [ESA]. I simply lay the matter before you ‘without bias.’” After appreciating visits to Wisconsin by the BAP’s A.D. (“Dog”) Middleton and Dennis Chitty, and by David Lack (a minor faux pas, as Lack, head of another Oxford zoology unit, and Elton were pursuing a personal “cold war,”) Leopold closed, 365 Meine, Aldo Leopold (cit. n. 163), 493. 165 adding a postscript: It seems needless to discuss the details of the proposed symposium until there is some reason to believe that you might participate.” Written by another hand on the archived carbon copy: “cc- [ESA Secretary, W.A.] Dreyer; [W.C.] Allee.”366 A second, more conspiratorial “Dear Charles” letter of the day was apparently dispatched together with the first: The attached ‘official’ letter discharges my official duty. I now wish to speak to you personally. What about a summer trip, bringing Jay [sic] along, and headquartering at our house, with no speeches? You could leave Jay with us during any side-trips you care to make for consultation purposes. You and Jay could also use our shack for as much ‘rustication’ as you have appetite for. Estella and I could join you there weekends. The difficulty, of course, is that such a venture would sacrifice the financial help of travel costs. I suppose it would be hard to get funds for a trip without speeches. Once over here, however, the trip would entail little outlay except for side-trips. The main purpose of this letter is to urge you to take a rest. If a summer trip headquartering with us has any appeal to you, Estella and I would be very glad, especially if Jay came too. I say ‘summer’ because up to May I am snowed under with courses and students. In late May I have promised the Park Service to go to Isle Royale to look over a problem of too-many-beaver and too-many-moose. Why not go with me? 367 Elton again replied quickly. This is a rare instance in which a letter from one man to the other and its specific reply were both preserved. He had evidently received the two invitations alluded to by Leopold as well. Likely irritated by Leopold’s reference to David Lack in the first letter, he was almost undoubtedly annoyed by the substitution of “Jay” for “Joy” in the second. In an 366 A. Leopold to C.S. Elton, 30 January 1947(a). ALP 9/25/10-3 001 002. 367 A. Leopold to C.S. Elton, 30 January 1947(b). ALP, Ibid. 166 awkward conflation of formal and informal regrets, Elton explained, “I think you will realise that if I were coming to the States, I should count it an enormous privilege to visit your Departments, and renew contact with my friends in Wisconsin. I certainly appreciate very warmly your invitation, and am sorry I cannot fulfill it. It is not only that I have a large number of matters to attend to in respect to my University and other scientific work (presumably everyone is in the same position there); but it is also that I genuinely require some time to think quietly about ecological questions and theories which I have only been able to do in odd times during the last six years. I am also pretty tired, and for this reason also wish to stay home and do some continuous and not too hectic research.” Elton added a postscript, by hand: “I hope this letter does not sound as if I had one foot in the grave; on the contrary. I want to do some steady work, and morale otherwise quite high also!” 368 Within a month, Elton sent another letter [not found] on behalf of Eugenio Morales-Agacino, a Spanish zoologist hoping to further his studies in Madison. Leopold replied with restrained optimism. “If he is not a candidate for a degree, but simply wants to come here and pick up the general atmosphere and take a few courses, the plan is feasible of execution, and I will be glad to do everything I can to carry it out. If on the other hand he is a candidate for a degree I would have to follow my present rule of turning down any and all comers. I now have eighteen graduate students and I am not doing justice to them. Beginning in 1948 it might become thinkable to take on special cases, since I hope by that 368 C.S. Elton to A. Leopold, 11 February 1947. ALP 9/25/10-3 001 002. 167 time to have some additional faculty.” A hand other than Leopold’s noted cryptically in the file copy’s margin, “Please destroy letter written to Dr Elton 3-2147 (it was not sent).”369 Leopold Formulates His Land Ethic; The Collaboration Ends “At some point, probably in late July” 1947, Aldo Leopold melded portions of earlier essays with new material to produce another piece, “The Land Ethic,” for his long-intended book of essays.370 He included a major normative claim, among the most often quoted passages of the entire Leopold canon. It usually appears somewhat out of context, and even readers already familiar with it as a sort of unrhymed couplet may not immediately recognize it as a paragraph: “The ‘key-log’ which must be moved to release the evolutionary process for an ethic is simply this: quit thinking about decent land use as solely an economic problem. Examine each question in terms of what is ethically and esthetically right, as well as what is economically expedient. A thing is right when it tends to preserve the integrity, stability and beauty of the biotic community. It is wrong when it tends otherwise.”371 Whether Elton saw this particular essay before its publication is not recorded. The array of values would have been familiar to Elton, but the logjam metaphor was iconic, American forestry. 369 A. Leopold to C.S. Elton, 28 March 1947. ALP 9/25/10-3 001 002. 370 Meine, Aldo Leopold (cit. n. 163), 501-505. At that point, Leopold’s working title for the collection was Great Possessions. 371 Leopold 1966 [1949], 240. True to his roots, Leopold chose a forester’s metaphor, the log that holds a floating logjam together, much as a keystone braces a masonry arch. 168 Also during the second half (and probably the third quarter) of 1947, Elton must have sent Leopold some documentation of the proposal spearheaded by Arthur Tansley (with Elton ‘in the van’) to reform, expand and bureaucratize British nature conservation.372 The plan, built in part on Elton’s wartime memo, was publicized in Tansley’s 1946 book Our Heritage of Wild Nature. Tansley’s closing comments were: The conservation of such areas is of three-fold importance. First there is their natural beauty and the deep sentiment which attaches to them as wild country in which one can escape from the strain of modern life; secondly there is their scientific value which has also essential economic implications; and thirdly their value as instruments of education in the widest sense. If we keep and manage them properly we can hand down to our descendants a Britain which is not only economically prosperous, but retains something of its distinctive natural historic character, continues to delight our sense of beauty, and affords rich material for the increase of knowledge. ‘Man shall not live by bread alone.”373 Leopold did not identify the documents he received, but he was overwhelmed by the extent of the plan, and doubtful of its prospects. He wrote to Elton, “I had no idea until I looked at the map how big a fraction of England is included in the proposed program. When we next have a chance to talk, which I hope will be soon, I will be intensely interested to hear whether the present government will have the courage to attack so large a task. Over here I see so many sound technical efforts stowed away on the library shelves that I have become skeptical.” Leopold informed Elton that he himself had finally agreed to fill in with a “discussion” at the December ESA meeting that Charles had earlier 372 373 A. Leopold to C.S. Elton, 8 November 1947. ALP 9/25/10-1 001 012. Arthur Tansley, Our Heritage of Wild Nature, (London: Readers Union/Cambridge University Press. 1946) 67. 169 declined to attend. Leopold also passed along the news that he had added a third member to his staff. “On your next trip over here I hope that both he and my faithful helper Bob McCabe can become acquainted with you.” There was also a wistful horticultural update: “The Elton oaks were girdled by mice again last winter, but one of them made a regrowth of 24 inches this summer, which is two inches greater than last year. I am beginning to think that in this world of mice and rabbits, an oak is a pretty old man before he even starts growing. You will remember that you planted these acorns ten years ago.”374 Leopold enclosed another “recent paper called ‘The Ecological Conscience.’” Flader and Callicott reported that this paper, [or a version of it] was delivered as a talk the previous June, and that “parts of it were revised and incorporated in ‘The Land Ethic’” then also in draft.375 The version presented by Flader and Callicott included, “The practice of conservation must spring from a conviction of what is ethically and esthetically right, as well as what is economically expedient.”376 On Wednesday, 14 April 1948, Aldo Leopold learned that the Oxford University Press would publish his book of essays.377 A week later he collapsed and died while fighting a fire on a neighbor’s property. Another week later, Robert McCabe distributed a letter describing the circumstances of Leopold’s death to “close friends,” the only Leopold-related correspondence preserved 374 A. Leopold to C.S. Elton 8 November 1947. ALP 9/25/10-1 001 012. 375 Flader & Callicott, River (cit. n. 182), 338. 376 Flader & Callicott, River (cit. n. 182), 345. 377 Meine, Aldo Leopold (cit. n. 163), 517. 378 among Elton’s personal papers at Oxford. 170 No condolences from Elton were found among the many archived in Leopold’s papers at the University of Wisconsin. Nearly nine years later, Charles Elton submitted a draft of the last in a series of three proposed radio talks to his BBC producer John Simons.379 As recorded, it included a statement strongly reminiscent of Leopold’s, if less concisely stated; and Elton’s was more like a set of premises than a conclusion: “I think of the human race as being on a very long train journey in company with all these [a million animal species] other passengers, and there seem to me to be three absolute questions that sit rather patiently waiting to be answered. The first, which isn’t usually put first, is really religious. There are some millions of people in the world who think that animals have a right to be left alone, or at any rate that they shouldn’t be persecuted or made extinct as species. Some people will believe this even when it’s quite dangerous to themselves. Efforts to control plague rats in some Indian warehouses have sometimes been frustrated because the man in charge put out water for the rats to drink. Ideas of this sort will seem folly to the practical Western man, or sentimental. Yet who can stand up and call them sentimental when a great scholar and prophet like Dr. [Albert] Schweitzer says: ‘The great fault of all ethics hitherto has been that they believed themselves to have to deal only with the relation of man to man?’ The second question can be called aesthetic and intellectual. You can say that nature—wild life of all kinds and its surroundings—is interesting, and usually exciting and beautiful as well. It’s a source of experience for poets and artists, of materials and pleasure for the naturalist and scientist. And of recreation. In all this the interest of human beings is decidedly put first. The third question is the practical one: land, crops, forests, water, sea fisheries, disease and the like. This third question seems to hang over the 378 R. McCabe to C.S. Elton, 21 April 1948. CSEP A.37. 379 C.S. Elton to J.R. Simons, 3 February 1957, BBC:ETF. 171 whole world so threateningly as rather to take the light out of the other two.380 Conclusions As stated at the outset, relevant documentation of the Leopold-Elton relationship is thin, and (presumably) woefully incomplete. Deductions based on it yield mainly hypotheses, not conclusions. But a few facts seem irrefutable. Early in their careers, before they met or influenced each other, Elton and Leopold each experienced and documented sentimental opposition to neobiota: Leopold in 1917, and Elton in 1925. Elton documented his concerns younger, but Leopold prior. Leopold’s, concerned with alien trout in the American southwest, were alarmist, buttressing a call to action. Elton’s, regarding squirrels in Oxford, were militarist, even in a letter to his mother; and in a published paper regarding blowflies in Spitsbergen; he adorned them with an openly anti-Roman Catholic remark. All these instances are merely the earliest available documents, and none seems likely to represent a real epiphany on the part of its author. At first glance, the conceptual similarities between Elton’s “three absolute questions” for the BBC audience and Leopold’s “key log” statement, the ethical/aesthetic/practical “tripod” on which each man rested his appeal for a new approach to conservation, supports David Takacs’ observation that Leopold’s conservation ideas influenced Elton’s.381 On the other hand, Tansley’s 1945 380 “The Conservation of Variety,” ‘as broadcast’ script BBC Written Archives B/C.T.P.30/3/57. 381 David Takacs, The Idea of Biodiversity. (Baltimore: Johns Hopkins, 1996), 22. 172 book, and perhaps the reports Elton forwarded to Leopold, prefigured both; but their seeds were already discernable in Elton’s 1942 conservation memo. For what it is worth, the University of Wisconsin Library owns a copy of Tansley’s book, and we know Leopold received a copy of Elton’s memo. There is no reason to argue otherwise than that, among these three formulations, Leopold’s was the most concise, most memorable and ultimately most influential piece of writing. It would be untenable on the basis of this study, and pointlessly provocative, to confiscate these ideas from Leopold and attribute them firmly elsewhere. But it might be wise to conceive of them as forged in the fires of a long conversation, rather than uniquely within Leopold’s personal reflections. By 1931, Elton was serving as something like Leopold’s ecological muse. Conversely, as time passed, Leopold inspired Elton’s conservation and outreach efforts. Perhaps also, as Leopold’s nature writing was eclipsing Elton’s in both quality and quantity, and as he (unknowingly) neared the end of his life, Aldo also assumed something of the ‘elder philosopher’ role brother Geoffrey had played during Charles’s youth. If Charles ever allowed himself to regard Aldo as a mentor in Geoffrey’s mold, he was doubtless nonplussed by their similarly sudden and unexpected exits. But as Leopold’s exit approached, Elton’s personal regard for him seemed to be waning. And a vague impression left by the matter-of-fact tone of Leopold’s last extant letter also suggests that he no longer really expected Elton to come back to America. And no suggestion was found that Charles and Joy ever invited 173 Aldo and Estella to Oxford, or for that matter that the Americans ever made plans, or even had a desire, to visit England. Epilogue Nearly twenty years after Leopold died, Elton recalled him as “the great natural philosopher of conservation in North America,” much as he acknowledged his own father Oliver as “a great literary critic” and John Muir as “the great American prophet of wilderness conservation.”382 But in a jarring and rather sad finale, Elton began his tribute, “In 1953, Aldo Leopold … wrote that ‘one of the penalties of an ecological education is that one lives alone in a world of wounds. Much of the damage inflicted on the land is quite invisible to laymen.’” This exclusionary contention is still beloved of many who claim such an education. Elton blunted its point, continuing, “As a matter of fact, much of it has been largely invisible to ecologists also, at any rate in this country.”383 More subtly, in mistaking the edition date of the Sand County Almanac on his shelf for the date of its authorship, Elton, carelessly or otherwise, overlooked the fact that by 1953 his late friend Leopold was no longer writing, and was in fact already five years gone. Six weeks after learning of Aldo Leopold’s end, Elton read another of the many unsolicited letters that constantly filtered into the BAP. This one arrived from South America. It commenced a relationship with a scientist whose outlook 382 Charles S. Elton, The Pattern of Animal Communities (New York: Wiley,1966), 382; The Ecology of Invasions by Animals and Plants (London, Methuen, 1958) 143, 159. 383 Elton, The Pattern of Animal Communities (cit. n. 382), 382. 174 and attitude toward neobiota would irritate Elton, and perhaps decisively nudge him toward writing his sixth and most fervently ideological book. Chapter 3: War and Peace: Views of Neobiota in the 1950s World War Two could be experienced as a soldier, or as a civilian. It could be experienced on or off the battlefield. It could be experienced as a preoccupation, or as an annoyance. Charles Elton had drilled with the Army during World War One, but was never deployed overseas. Anticipating that one way to stay off the battlefield was to make some other, useful contribution, he had enlisted the BAP for rearguard service against rodent depredations. This postponed all other BAP research. Furthermore, he was in a country that, if not a battleground in the same sense as continental Europe, was certainly on the sending and receiving ends of a great deal of ordnance. Elton was preoccupied by the war effort. American Marston Bates, a Harvard-trained tropical field ecologist, was working in Egypt on mosquito-borne diseases for the Rockefeller Foundation when the war broke out in Europe. By the time Pearl Harbor was attacked, he had established a new research base to Colombia. Bates was dislocated by the war, but not preoccupied by the war effort. In late 1954, Bates began preparing a review of neobiotic phenomena. He developed it as a pre-circulated symposium paper that was subsequently published in a widely available book of proceedings. Oxford animal ecologist Charles Elton lukewarmly acknowledged that publication in the preface to his own (1958) The Ecology of Invasions by Animals and Plants: “I have found useful 176 references in a paper by Marston Bates (1956), ‘Man as an Agent in the Spread of Organisms’…this is the only recent general review of the subject of invasions that I have seen.”384 After The Ecology of Invasions appeared, “Man as an Agent,” was nearly forgotten, despite two years’ publication priority and a contextually more “scientific” advent compared to Elton’s book. This chapter introduces Bates, briefly compares his career with Elton’s, and describes the genesis of each man’s monograph on neobiota. It demonstrates the existence of a substantial alternative to Elton’s militarized conception of ecological “invasions,” demonstrating that warfare was not the only possible, inevitable, or necessary ecological model of neobiota at the time. Finally, it proposes some reasons for the differential persistence of the two authors in the institutional memory of invasion biology. Natures and Nurturings Marston Bates had three careers; first as a research entomologist, second as a university professor, and third, bridging the first two, as a writer of popular books and commentaries on natural history.385 Born in Michigan but raised in Florida, he was an instinctive adventurer, comfortably adapting to living and working in culturally unfamiliar settings including Albania and the ecologically riotous American tropics. 384 385 Elton, The Ecology of Invasions (cit. n. 162), 7. Bates was Stephen Jay Gould’s predecessor as a regularly featured columnist in Natural History magazine. 177 At the beginning of the twentieth century, Marston’s father Glenn Bates worked in the exotic plant trade in south Florida.386 He was befriended there by David Fairchild, son-in-law of Alexander Graham Bell, and an avid collector and promoter of plants who credibly claimed general responsibility for importing to the United States some 200,000 species and varieties.387 Fairchild’s son Sandy was the same age as Marston, and both were armed with butterfly nets, leading to a long association.388 In 1934, after Marston completed his Harvard Ph.D. in zoology he returned to Florida, briefly courted Sandy’s sister Nancy Bell, married her under a Javanese fig tree at the Fairchild estate, and then whisked her away to a “an old Turkish house with its floor made of mud mixed with ox blood” in Albania, where he studied malaria transmission for the Rockefeller Foundation.389 Bates’s malaria research was interrupted and relocated by the outbreak of World War II. Marston and Nancy moved briefly to Egypt and thence to Colombia. There, for eight years, he studied yellow fever’s mosquito vectors and pioneered the practice of erecting platforms at various levels of the forest canopy 386 William C. Kimler, “Bates, Marston.” Dictionary of Scientific Biography (New York: F.L. Holmes, 1981), 17: 51-53. 387 David Fairchild, Exploring for Plants (New York: MacMillan, 1930); David Fairchild, Elizabeth Kay and Alfred Kay. The World was My Garden: Travels of a Plant Explorer (New York, Charles Scribner’s Sons, 1938). 388 Nancy B. Bates, East of the Andes and West of Nowhere, (New York: Scribner’s, 1947), 16. 389 Ibid,.16-17; David Fairchild, The World Grows Round My Door (New York: Scribner's, 1947), 314. 178 in order to study insect habitat stratification.390 His in-laws were comfortably well off as a result of Bell’s telephone patents, and were regular benefactors of, and (sometimes extended) visitors to, the Bates home in South America.391 In 1948 the Bates family left Colombia for a New York administrative assignment with the Rockefeller Foundation. Then, in 1952, Marston accepted a Professorship in Zoology at the University of Michigan. Arriving in the semi-tolerable climate of Ann Arbor, Bates converted a disused porch of his college town residence into a heated greenhouse, stocking it with tropical plants, finches and hummingbirds, lizards, insects, and even monkeys.392 At age 58, when visions of a secure retirement loom in many minds, he reminisced wistfully about the contingencies of rural Colombia: “We had a whole range of tropical landscapes within easy reach of our town– only desert and seashore were lacking. This was fascinating enough in itself, but beyond us there was, ever present, the thrill of possible encounter with the unknown.”393 In the atmosphere of a liberal American college town, he was gregarious to the 390 Marston Bates, The Forest and the Sea, (New York: Random House, 1960) 18. 391 David Fairchild, Garden Islands of the Great East (New York: Scribner’s, 1944); N. Bates, East of the Andes. 392 393 Marston Bates, A Jungle in the House, (New York: Walker and Co, 1970). Marston Bates and the Editors of Life Magazine, The Land and Wildlife of South America (New York: Time Inc.), 9. 179 point of presiding (in his own home) over an almost daily group dinner and discussion session attended by his own students and other hangers-on.394 Contrasting sharply with Bates, Charles Elton was born into a family of literati. His “upbringing had many aspects that were characteristic of comfortable middle-class Edwardian families: nursemaids—mostly harsh—preparatory school and bullying, public school, army cadets, [and] long summer holidays when the whole family moved to the country.”395 Charles admittedly owed his interest in natural history to his acute but luckless eldest brother Geoffrey, who instructed, encouraged and cajoled his younger sibling until his own sudden death at age thirty-three.396 Elton was a career Oxford man. His student fieldwork took him repeatedly to the biologically depauperate Arctic, and he conducted most of his subsequent research in Wytham Woods, a few miles from his University office. Although he long presided over the Oxford University Exploration Club, he rarely left England, after 1938, and visited the tropics only after his retirement. As an adult he was diffident, even semi-reclusive; “he avoided the public occasion.”397 Elton’s student (and subsequent Bureau staff member) Kitty Paviour-Smith described 394 Personal communication: James Collins, Arizona State University, May 2004; Doug Futuyma, University of Michigan, June 2004. 395 Southwood and Clarke “Charles Sutherland Elton” (cit. n. 211). 396 Charles S. Elton, unpublished ms. CSEP: A19, A20, A21, A33, A34. 397 Southwood and Clarke, “Charles Sutherland Elton.” (cit. n. 211). 180 him as “never an easy person to know.”398 He lectured only to make ends meet, having “neither the wish nor the aptitude to undertake any college teaching.”399 Nevertheless Elton founded and directed the Oxford Bureau of Animal Population (BAP), and managed to find enough “soft money” research funding from businesses, governments and organized interest groups to keep it afloat for thirty-five years.400 He too presided over a daily gathering: afternoon tea in the BAP library.401 If these brief sketches adequately suggest their worldviews, or at least their world preferences, perhaps one could conclude that Elton had the more urgent and conservative sense of propriety and yearning for stability. It is tempting to shorthand that by simply stating that Elton was an Englishman and Bates, an American. But they had something in common. Each in his own way had a commanding presence, and each, within his circle, was commonly addressed as “The Boss.” 402 398 Kitty Paviour-Smith, “Elton, Charles Sutherland (1900-1991)” Oxford Dictionary of National Biography (Oxford University Press, 2004) online at http://0-www.oxforddnb.com.library.lib.asu.edu:80/view/article/49632, accessed 19 July 2005. 399 Southwood and Clarke “Charles Sutherland Elton.” (cit. n. 211). 400 Peter Crowcroft, Elton’s Ecologists (cit. n. 184). 401 Ibid., 148. 402 E.g., Ibid., 67; Nancy Bates, East of the Andes (cit. n. 388), throughout. 181 The Nativity of “Man as an Agent” The published review paper “Man as an Agent In the Spread of Organisms” was one among an array of contributions Bates made to a 1955 international symposium titled Man’s Role in Changing the Face of the Earth.403 Understanding the symposium is helpful to gauging the significance of Bates’s offering. The meeting was sponsored by the Wenner-Gren Foundation for Anthropological Research, and held in Princeton, New Jersey. Bates was a coorganizer and co-chair of the meeting, responsible for the general division on “process” (ongoing change, as distinct from the “retrospect” and “prospect” divisions which were respectively chaired by geographer Carl O. Sauer and critichistorian Lewis Mumford). Bates also chaired a session called “Changes in Biological Phenomena” and helped edit the exceptionally detailed, nearly 1200page symposium proceedings volume underwritten by the U.S. National Science Foundation.404 After nearly three years of planning, entailing many metamorphoses, the Man’s Role symposium was formally announced in three major and laconically 403 Marston Bates. “Man as an Agent in the Spread of Organisms” IN William Thomas, et al, Man’s Role in Changing the Face of the Earth (Chicago: University of Chicago Press. 1956) 788-804. 404 William Thomas, “Introductory” in Thomas, et al, Man’s Role in Changing the Face of the Earth, (see cit. n. 403) xxi-xxxviii. 182 titled journals: Man, Science, and Nature.405 Seventy notables representing fifteen countries participated. Seventeen spaces were specifically reserved for (and taken by) scholars from “outside the United States,” and eight of those seventeen were British, but none were Oxford scholars. Among those representing Britain were Leslie Banks, Cambridge Professor of Human Ecology; Fraser Darling, Senior Lecturer in Ecology and Conservation at the University of Edinburgh and Sir Charles Galton Darwin, with pedigree, but without portfolio. All participants were provided with first-class travel and accommodations.406 At least one recent “retrospective reviewer” considered the meeting to have been disappointingly short on activism. In the July, 2005 Environmental History, Syracuse University Geographer Robert M. Wilson concluded: “[A]s a geographer looking at Man’s Role in Changing the Face of the Earth a halfcentury later, one can only be dismayed that geography did not seize an opportunity to serve as one of the intellectual cornerstones of the environmental movement during its formative years.”407 However, this criticism seems both anti-historical and anachronistic. The organizers plainly stated their intentions in advance: 405 In his “Introductory,” Thomas wrote that the “first public announcement” appeared in the March 11, 1955 issue of Science (121:356-357), but it actually appeared some seven months earlier under a group byline in Man (54:139). The Nature (175:754) announcement came in late April, 1955. It was shortened by about half with each successive publication. 406 407 Thomas, “Introductory.” (cit. n. 404) R.M. Wilson, “Retrospective Review: ‘Man’s Role in Changing the Face of the Earth’ Environmental History 10 (2005), 564-566. 183 The symposium idea presents an important intellectual problem with a focus of interest for persons with different theoretical and descriptive backgrounds. Here, we are trying for an overview of the state of knowledge on a topic touched upon only in piecemeal fashion by individual disciplines. Emphasis is to be placed on the stimulation of cross-disciplinary thought, a review of what is not yet known, and the delineation of the most profitable lines of future research. No resolutions, no formulation of an action programme, no incorporation of an organization are contemplated. This is intended as a gathering of some 80 of the world’s best minds in a congenial atmosphere for recorded discussion leading to eventual publication for the benefit of the world of scholarship.408 The earliest extant “Tentative Program” among the symposium documentation in Bates’s papers, dated 3 February 1954, was titled Man’s Impact rather than Man’s Role. A proposed Thursday (day four) session called “Man’s Effects on Plants and Animals,” to be chaired by palynologist Paul Sears, included a paper by Bates: “Man’s Competition with Plants and Animals,” and another by Frank Egler; “Effects of Intentional and Unintentional Transport by Man.” Egler, then a research associate at the American Museum of Natural History’s Department of Conservation and General Ecology, showed his interest in species introductions with the 1942 Ecology paper "Indigene Versus Alien in the Development of Arid Hawaiian Vegetation" and several later publications.409 This division of labor between Bates and Egler survived into a later (but undated) agenda carrying the ultimate symposium and session titles, but was not realized in the symposium itself. 408 Carl Sauer,et al., “An International Symposium: Man’s Role in Changing the Face of the Earth” Man 54 (1954),139. 409 Frank E. Egler. “Indigene vs. Alien in the Development of the Arid Hawaiian Vegetation.” Ecology 23 (1942), 14-23. 184 Only fragmentary documentation of what transpired next survives among Bates’s papers.410 Egler attended the meeting, but evidently begged off writing duties. Abhorring a topical vacuum, Bates scrambled to fill it with a review of his own. A June 1954 pro-forma symposium invitation to Bates from the WennerGren Institute identified him as the prospective author of a paper “on the subject of ‘Plant and Animal Diseases and Pests Spread by Man’… to be submitted not later than March 15, 1955.”411 Shortly thereafter, Bates wrote to Wenner-Gren’s William Thomas, “I want to start thinking about the material for my own paper. I have been wondering about a change in the title-- ‘pests’ is not one of my favorite words. What about ‘Man as an Agent in the Spread of Organisms’ or the ‘Effects of Man on Biogeography’ or something of that sort. It seems fairly clear that what you are after is a summary of man as an agent of distribution change on a geographic scale.”412 Bates prevailed. In early December 1954 he wrote to Egler, “I’ve started thrashing around trying to get my paper organized -- ‘Man as an agent of distribution of organisms.’ My ignorance of the literature is vast. Is there any general survey of the accidental and purposeful plant introduction into Hawaii? Have you any general suggestions about what I should deal with and where to 410 Frank Egler’s papers have been collected, but not catalogued, at his Aton Forest, Connecticut research site. 411 George Fejos to M. Bates, 21 June 1954. Marston Bates Papers [MBP], 6-3 University of Michigan Bentley Historical Library. 412 M. Bates to W. Thomas 13 July 1954. MBP 6-3. 185 start looking it up?’ Bates closed his query, “Don’t go to any trouble-- I’m just betting you’ve got some good ideas at your fingertips.”413 Even if his ignorance of the literature was vast, Bates at least had recent, real-world experience with the problem of categorizing plants and animals according to their origins. From late June to early September 1953 he had participated as the “land biologist” for an interdisciplinary team of Pacific Science Board researchers surveying Ifaluk Atoll in the Caroline Islands, two days’ sailing south of Guam.414 One of his objectives was to census terrestrial species, while one general goal of the project was anthropological study of the islanders. Bates described the problem of this interface as follows: Low-lying Ifaluk has probably been a completely submerged reef within a matter of a few thousands of years. All of the plants and animals living on Ifaluk, then, have had to have some means of getting there within a comprehensible period of time. The plants and animals, from this point of view, fall easily into three groups: those that got to Ifaluk by some 'natural' means, those deliberately brought by man, and those accidentally introduced by man. I tried to sort out the 119 species of plants according to these three categories, but it turned out not to be easy. Thirty-two species were clearly always cultivated, and were equally clearly deliberate introductions by man. But there are many borderline cases [e.g. coconut palms]. It is also, in many cases, hard to distinguish between accidental human dispersal and natural dispersal. Many plants have natural means of dispersal that enables them to get almost everywhere … [b]ut it is beyond my power to make a clear decision in every case. Roughly, I suspect that about a third of the plants of Ifaluk got there by natural means– would be there if man 413 414 M. Bates to Frank Egler, 8 December 1954. MBP 6-24. Marston Bates and Donald Abbott, Ifaluk, the Story of a Coral Atoll. (London: Museum Press 1959) 5-19. Ifaluk is now part of the State of Yap in the Federated States of Micronesia. 186 had never taken to navigating– and that another third were brought by man deliberately, and the remaining third by man accidentally.415 Egler responded hesitantly to the appeal for assistance, writing “that is by no means my forte,” but offering several suggestions for reading and personal contact. He also offered skepticism. “As an Auslander, [to plant geography] I am all confused from the literature as to how much imagination, as well as ‘real’ taxonomy is involved in their currents, fruit-eating birds, land bridges, and hypothetical early migrations of man. Everyone has everything going in different directions by different means; and it all sounds equally sensible.” Enclosing a reprint of his twelve-year old Hawaii paper, he added, “I still have the same opinions– more so; but I know it is a very irreverent attitude, in contrast to those men who like to fear the aliens.”416 Some of those opinions looked like this: When the author arrived in Oahu in 1936, he soon became conscious of a strong antipathy among certain laymen towards the naturalized foreign plants. How such an antipathy can actually discolor one's interpretation of existing conditions was exemplified by a student who claimed that the alien flora apparently possessed a certain group spirit and cooperative action which permitted it to carry on a mass "warfare" against the indigenous flora.417 Not all mature and serious considerations of the vegetation problem in Hawaii, contrary to the case mentioned above, assume that aliens are replacing indigenes by actual competitive superiority. The thesis which does dominate the literature is that the original equilibrium has been disturbed by man, fire and grazing animals, succeeding which, alien plants have invaded and established a new and more or less permanent "balance of nature" that rigidly excludes the indigenes. An undercurrent of 415 Bates and Abbott, Ifaluk,119-120. Ifaluk was published in 1958 in the United States under the title Coral Island. There is no evidence that Bates read Elton’s The Ecology of Invasions before Ifaluk went to press. 416 Frank Egler to M. Bates, 10 December 1954. MBP 6-24. 417 Egler, “Indigene” (cit. n. 409) 15. 187 aversion towards the alien flora and a minor tone of gloom for the future of the remaining indigenes are usually present. [Italics in original.]418 Three months later (and a week ahead of deadline), Bates submitted his Egler-influenced paper to Thomas, noting in a cover letter “It seems to me only a beginning of a review of the subject, but I take comfort in thinking I’ll have a chance to write it over again before publishing.”419 It is unclear whether he ever did so. The only prepublication version of “Man as an Agent” surviving among Bates’s papers is an undated, hand-corrected typescript filed separately from the letter to Thomas, resembling the published version in all but one significant respect: it included a preamble that was dropped from the published draft.420 Like the other symposium papers, “Man as an Agent” was distributed for advance reading rather than presented from the podium; the sessions themselves consisted of conversation based on the written materials. The published (edited) record of the session discussions makes no specific mention of Bates’s paper or topic, but it does summarize substantial remarks by Egler and several others (among them, John T. Curtis, Clarence Glacken, and Edward H. Graham) regarding then-current concepts of ecological equilibrium.421 418 Ibid., 16. 419 M. Bates to W. Thomas, 7 March 1955. MBP 8-21. 420 Marston Bates, Undated typescript: “Man as an Agent in the Spread of Organisms” MBP 19-5. 421 Arriving at the BAP for the first time as a student, Peter Crowcroft asked Charles Elton for a reading list. Elton replied “You had better start by reading Ed Graham’s Natural Principles of Land Use.” (Elton’s Ecologists 141); but Graham’s work is not particularly well remembered today. 188 The published symposium proceedings did not go unnoticed in Britain. Max Nicholson, Director-General of Britain’s (official) Nature Conservancy and Elton’s colleague on the Wild Life Special Committee reviewed Man’s Role in some detail for the November 1957 issue of The Journal of Ecology. He specifically mentioned that “Dr. Marston Bates in Man as an Agent for [sic] the Spread of Organisms reviews deliberate and accidental introductions of animals and plants, and finds that easy generalizations do not fit and the ‘experiment-like situations produced by man’s alteration of environmental factors, and by his movement of organisms into different environments have not been fully appreciated.”422 The BBC Proposes and Elton Disposes In late September or early October 1956, the British Broadcasting Corporation (BBC) contacted Charles Elton, requesting that he prepare and present a series of radio talks on The Third Programme, (Programme, below) which was directed at “persons of taste, of intelligence, and of education.”423 This was not an unprecedented commission, but the proximate motivation for the BBC’s interest in Elton at that time is unclear. Over the years many scientists had gone “on the air” for Britain’s state broadcasting service, Elton included—in 1933 and 1937. Except for his election as a Fellow of the Royal Society in 1953 and more recent but publicly uncontroversial (even unnoticeable) resignation 422 E.M. Nicholson, (Review of ) Thomas, William L. Jr. (Editor) “Man’s Role in Changing the Face of the Earth.” The Journal of Ecology 45 (1957), 955-958. 423 Humphrey Carpenter The Envy of the World: Fifty Years of the BBC Third Programme and Radio 3 (London: Phoenix Giant 1997), 12. 189 from the Nature Conservancy’s Scientific Policy Committee, Elton was not very newsworthy. Neither was he a charismatic public intellectual like Julian Huxley. To most of his potential listeners, he was practically a cipher. A BBC staffer tried “for some time to bring Charles Elton to the point where he would offer us a talk,” describing the ecologist as “a shy bird when it comes to public appearances.”424 This observation was echoed by an Elton acquaintance: “he could not abide crowds, and refused to attend meetings with more than thirty people present.”425 The thought of speaking to an audience of thousands, even unseen, doubtless made him uncomfortable. Perhaps this was mitigated by the fact that the series would be recorded for later transmission. The choice of topic was more or less left to Elton’s discretion, and if he considered various options there seems to be no record of those he rejected. In late October, Elton submitted a draft of one talk and outlines of two more. These documents did not survive in the BBC archives, but their contents produced guardedly optimistic reactions. Staffers wrote that the draft “becomes very much a lecture after page four, but I think the theme he has chosen, particularly for the last talk, is very fruitful,” and “As it stands, this seems a bit undisciplined … but it is good lively stuff & seems to promise the right kind of talk.”426 It appears that a meeting between Elton and two or more BBC 424 J.C. Thornton to C. Holme, 5 October 1956. BBC:ETF. 425 Kitty Paviour-Smith, “Elton, Charles Sutherland (1900-1991).” (cit. n. 398). 426 J.C. Thornton to A. Clow, 23 October 1956; [unidentified, handwritten note] to P.H. Newby 31 October 1956. BBC:ETF. 190 personnel occurred shortly thereafter. But some concerns remained. A month later, P.H. Newby, who would have a distinguished career at the BBC, wrote in a memo: “see [Elton] as soon as possible. I should like to have a word with you about tactics, because I think that Dr. [sic] Elton needs careful handling.”427 All apparently went well enough. On 18 January 1957, the BBC sent Elton a brief formal invitation offering him sixty guineas and transportation costs. He worked quickly, submitting full drafts of the first two talks to his producer John Simons a week later, with the third following on 3 February. The talks were recorded in London on 20 February and 1 March 1957, for broadcast later in March under the collective title “Balance and Barrier.”428 Audience reactions to the first two broadcasts were mixed, but mostly positive. John Simons wrote to Elton, “You will be glad to know that your three talks were very well received by the general public, the first and third more so than the second … each of [them] received an appreciation index considerably above the average for the Third Programme, which for first broadcasts is a very pleasing thing.”429 The last comment might not particularly have pleased Elton, given his broadcasts in 1933 and 1937, of which Simons was apparently unaware. One bit of evidence suggesting that his message did not strongly resonate with the public is that there were no published letters to the editor of The Listener 427 P.H. Newby to J.R. Simons 27 Nov 1956. BBC:ETF. 428 Ronald Boswell to C. Elton, 18 January 1957. BBC:ETF. 429 J. Simons to C. Elton, 13 May 1957. BBC:ETF. 191 regarding Elton’s radio talks. BBC Audience Research Reports survive for the first two episodes; they confirm Simons’ interpretation in that both were rated as slightly above average in topical interest. Surveyed listeners appreciated Elton’s wit; but it is not recorded that any responded as if they had been called to action.430 Despite Simons’ assertion (and as suggested above) Third Programme broadcasts were not exactly meant to appeal to “the general public.” At just about the time that Elton was negotiating and preparing his talks, the Programme itself was coming under increasing scrutiny. In January 1957 a “working party” reported to the BBC Governors that it was “‘too much occupied with minorities within a minority’ and the use of such epithets as ‘precious’, ‘esoteric’ and ‘donnish’ had ‘some measure of justification.’ Too many people had been ‘put off by the general character of the Programme and … made to feel that [it] is too difficult and too high-brow and altogether outside their reach.’”431 Influence and Effect The crux of this particular comparison between Bates and Elton is that the two ecologists interacted while developing their respective 1950s publications. Like many others, Bates initiated contact with Elton as a supplicant. In 1945 Elton’s Journal of Animal Ecology published a paper by Bates on Colombian 430 BBC Audience Research Reports, 12 and 15 April 1957. BBC Written Archives R9/6/63; John Chrysostom, Homily 24 on the Epistle to the Hebrews. 431 Carpenter, The Envy of the World, (cit. n. 423) 169-170. 192 mosquito ecology.432 In mid-1948, still working in Colombia, Bates sent a letter requesting permission to employ Eltonian epigraphs for several chapters of his own first book The Natural History of Mosquitoes, which cited his 1945 paper and used some of the same photographs. Perhaps he charmed the Englishman by writing, “I seem to have picked on you more often than on anyone else with Lewis Carroll a close second.”433 Elton graciously acceded to the request, extolling Carroll and claiming him as a fellow Oxfordian.434 (Disingenuously, it seems; Elton disclosed elsewhere that he considered Carroll “cruel” and “artificial”).435 Hard on the heels of Bates’s first book came a second, The Nature of Natural History, crediting Elton with many ecological conceptions and discoveries (via his books Animal Ecology and Voles, Mice and Lemmings). In mid-October of that year (1950) Bates visited the Bureau of Animal Population for a few days “to discuss human ecology problems,” an event duly noted but not elaborated in Elton’s “Diary of Events, 1949-1952.”436 The two authors at least occasionally exchanged books; their last surviving correspondence consists of a transmittal note from Elton accompanying a copy of his final (1967) volume The Pattern of 432 Marston Bates, “Observations on Climate and Seasonal Distribution of Mosquitoes in Eastern Colombia” Journal of Animal Ecology 14 (1945) 17-25. 433 M. Bates to C.S. Elton, 3 June 1948 MBP 4-36. 434 C.S. Elton to M. Bates, 14 June 1948 MBP 4-36. 435 C.S. Elton, unpublished ms., CSEP A33:10. 436 Crowcroft, Elton’s Ecologists (cit. n. 184), 158. 193 Animal Communities, and Bates’s acknowledgement thereof.437 Copies of several of Bates’s books reside in Oxford University’s Elton Library, at least one bearing the inscription “presented by the author.” There are three hints that Elton did not quite view Bates as a colleague of equal standing. The first was his dismissal of “Man as an Agent” as a source of useful sources for his own book, without citing the paper itself as a significant precursor. The second came in Rick Miller’s reply to Elton’s June 1957 letter about the BBC assignment: “I was tremendously impressed by the Third Program [sic] series… you’ll be quoted and quoted— had Marston Bates had access to it, Lewis Carroll wouldn’t have had a chance in The Nature of Natural History!” Noting but otherwise ignoring this invocation of the wrong Bates title, it is interesting that the Carroll episode remained somehow relevant nine years after the original fact. Furthermore, it was mentioned not by Elton, but by Miller, who arrived in Oxford sixteen months behind Bates’s 1948 letter. The anecdote must have been a favorite of Elton’s, at least during Miller’s Oxford tenure (1949-1951) or it would have faded from consciousness long before. Why it was a favorite remains a matter for conjecture. Most intriguingly, in February1955, just as Bates was overcoming his “vast” ignorance and thrashing out a thesis based on Egler’s referrals, Elton wrote to enlist Bates’s advice and help in securing a tropical forest ecology position for E.W. Fager, an Oxford doctoral student. Bates dutifully forwarded 437 C.S. Elton to M. Bates, 13 June 1965; M. Bates to C.S. Elton, 26 July 1966. MBP 11-2. 194 the request to his contacts at the Rockefeller Foundation and Smithsonian Institution, advising Elton of this action and offering a few relevant observations.438 He closed by musing: I have got off into human ecology and seem to be stuck there—but it is a fascinating business. I spent several months a while ago on a small and remote atoll in the Pacific, trying to get some measure of the relations between man and the environment in this supposedly ‘simple’ situation. Clearly, there, man was the ecological dominant really responsible for the total vegetation pattern of the island. More and more I have been impressed by the failure of our ecologists to take adequate account of man as an ecological agent. They seem often to pretend that he doesn’t exist when he has really completely made over the landscape.439 A few days later, Elton responded with thanks, and musings of his own: Your coral atoll survey sounds very attractive. Do you remember about the natives who explained the arrival of human fleas on their islands by the notion that they were the restless souls of white men? I suppose there are absolutely no islands left which have not been subject to the shocks of outside visitors. It must be a change for them to have someone like you who is interested in their whole ecology. In establishing National Nature Reserves in this country, through the [governmental] Nature Conservancy, we have run into very interesting examples of the unsuspected influence of man.440 Elton followed the last remark with several examples, none of which had to do with neobiota. Bates mentioned fleas in “Man as an Agent,” but only as meeting the criteria of being both inquilines (a conception he fancied, and promoted elsewhere), and as parasites.441 438 C.S. Elton to M. Bates, 9 February 1955. MBP 6-24. 439 M. Bates to C.S. Elton 15 February 1955. MBP 6-24. 440 C.S. Elton to M. Bates, 28 February 1955. MBP 6-24. 441 M. Bates “Man as an Agent,” (cit. n. 403) 795. “Inquiline” refers to a form of non-injurious symbiosis involving an animal that lives in the nest or home of another species. 195 Elton, of course, included fleas in The Ecology of Invasions, where they closed out the chapter “The Fate of Remote Islands”: “Perhaps we could bear in mind the story told by Buxton and Hopkins, about the arrival of the human flea in one small Pacific Island [sic] more than a hundred years ago: ‘The placid natives of Aitutaki, observing that the little creatures were constantly restless and inquisitive, and even at times irritating, drew the reasonable inference that they were the souls of deceased white men.’ We may hope that this same restless curiosity in the form of research will find out how the broken balance can be restored and protected.”442 Questions hover around this episode, but, “that’s all they wrote,” or at least all they left us to ponder. Why didn’t Bates more explicitly ask Elton for ideas or examples? Why didn’t Elton mention his own previous work, instead of referring to a cryptic and (in the letter) fragmentary anecdote? Had he related it to Bates at Oxford in 1950? One preliminary objective of this study was to determine whether Elton used the radio talks as a basis for his 1958 book, or (as might be likely a few decades later) as a way to promote it. Almost three months after the recording sessions, Elton wrote to his sometime collaborator and former post-doctoral research assistant Richard S. (Rick) Miller, “[I] did the broadcasts I sent you copies of; am now turning that stuff into a 45000 word heavily illustrated book for Methuens which I shall have written in nine weeks, ten days more to go, mostly 442 Elton, Invasions, (cit. n. 162) 93. 196 about invasions and ecological interspersion, etc.”443 Seven months later, he finished the story in another letter: “I got the book finished by the end of July … I got the writing done in three months … Of course I had the broadcasts as a basis, and much of the material already indexed over the years; nevertheless I did quite a blitz on further reading.”444 The best available chicken-or-egg answer appears to be that BBC’s request ultimately precipitated the book’s particular formulation. Based on his letters to Rick Miller, about a month elapsed between the second and final recording date and the beginning of earnest work on the book. As shown in Chapter one, Elton had the topic of neobiota on his mind for decades prior; but it cannot yet be determined with certainty whether he was specifically contemplating a book project in advance of his BBC commission. As archived by the BBC and published in The Listener, the “Balance and Barrier” transcripts more or less resemble four chapters (one, six and a combined eight-and-nine) of the ensuing book. On the air, Elton mentioned fewer cases and employed a chattier vernacular, but the book was also intended for an educated, non-specialist audience and produced by an all-purpose British trade publisher. A comparison of the radio and book versions indicates that Elton continued to refine and occasionally significantly revise his ideas as the book took shape. The exercise is appealing, but a full analysis of the ninety-day metamorphosis from scripts to book would require an extended discussion 443 C.S. Elton to R.S. Miller, May 26 1957. CSEP E.28. 444 C.S. Elton to R.S. Miller, January 30 1958. CSEP E.29. 197 without contributing substantially to the objectives of the present project, and must be undertaken elsewhere. Did Bates, with this correspondence and/or his symposium paper, significantly rekindle or refocus Elton’s interest in neobiota? Elton submitted his first draft of “The Invaders” to the BBC in mid October 1956, the year Man’s Role in Changing the Face of the Earth was released in print. There is a copy of the symposium proceedings in the Oxford Zoology Library, which incorporates the holdings of Elton’s Bureau of Animal Population, but no record or notation of its accession date.445 We cannot know with certainty whether Elton read “Man as an Agent” before he agreed to the BBC project. But he had obviously read it before prefacing his ensuing book. Max Nicholson’s November 1957 review of Man’s Role more or less marks the last possible moment for Elton to become aware of Bates’s paper. If Elton really finished writing in July 1957 as he reported to Rick Miller (recalling the apocryphal eighty-five day composition of Animal Ecology, and that the letter to Miller was dated January 1958), the published Nicholson review would appear to have no obvious bearing on the issue.446 However, Nicholson was a longtime acquaintance of Elton’s, and would have been well aware of Elton’s interest neobiota. The Journal of Ecology published no “books received” notices, so we do not know when Nicholson began reading. But it presumably took some time 445 Linda Birch, Oxford Zoology Department Librarian to M. Chew, 4 January 2005. 446 Elton’s “Preface” to Invasions (cit. n. 162), mentioning Bates, was dated 24 July 1957. 198 to digest the 1,100 pages of Man’s Role for a review, and he could easily have dropped a “heads up” note to Elton within the requisite period. It is conjectural, because no evidence of such a communication was found among Elton’s papers; but that is a very incomplete archive. Taking another tack, of the eight references the two publications share, only two are cited in the radio-derived chapters (one, six or eight-and-nine) of The Ecology of Invasions. Hard evidence that Elton was unfamiliar with either of these before “Man as an Agent” was published might provide the answer. One of them was Anopheles gambiae in Brazil, 1930-1940 by Fred Soper and Bruce Wilson, published in 1943 by the Rockefeller Foundation (a longtime Bates employer). This volume resides in multiple Oxford libraries, and Elton reviewed it in 1944 for the Journal of Animal Ecology (see Chapter Two). The other was The Dispersal of Plants Throughout the World, by Henry N. Ridley, tropical botanist, former Director of the Singapore Botanic Garden, and (nearly sixty years prior) an acquaintance of Bates’ father-in-law, David Fairchild.447 Several copies of Ridley’s book reside in library stacks at Oxford, as well they should, and probably have since its 1930 publication. It was reviewed in February 1931 for The Journal of Ecology; like Man’s Role, Dispersal could have come to Elton’s attention that way. Elton’s only published description of plant dispersal during that era came in his 1933 BBC talks and book, Exploring 447 Fairchild, Garden Islands (cit. n. 391) 84-85. Fairchild called Ridley “an outstanding scientist” whose “discovery of the correct method of tapping the Brazilian rubber trees was the turning point in the cultivation of rubber on a plantation basis in the Orient.” 199 the Animal World (see Chapter Two). Neither version mentioned sources, but Ridley’s book certainly did not account for the anecdotes Elton related to his 1933 listeners and readers. So it remains plausible, and perhaps even likely that “Man as an Agent” actually helped motivate Elton’s The Ecology of Invasions, but no “smoking gun” evidence or inevitable circumstance confirms it. Two Views of Neobiota It was Elton’s longstanding opinion that neobiota amounted to anthropogenic ecological war. Bates included a few allusions in this direction, but whether he was feeling somewhat bound to objectivity by the context in which he spoke (especially as a meeting organizer), was less convinced (like Egler) that he was dealing with a major problem, or was reluctant for some other reason, he did not issue a call to action or intervention. As Nicholson had pointed out, Bates described man’s agency in the spread of organisms as difficult to generalize about, but nevertheless generating “experiment like conditions” under which to study ecology, offering “great possibilities for the understanding not only of ourselves but of the system of nature to which we belong.”448 Perhaps his most judgmental remark was: As biologists, we are apt to deplore [the pervasive influence of humans], to brush it off, to try to concentrate on the study of nature as it might be if man were not messing it up. The realization that, in trying to study the effect of man in dispersing other organisms, I was really studying one aspect of the human habitat came as a surprise to me. But, with the realization clear in my mind, I wonder 448 Bates, “Man as an Agent,” (cit. n. 403) 802. 200 why we do not put more biological effort directly into the study of this pervasive human habitat.449 Numerous reports of neobiota had come to light by 1954. Bates mentioned at least fifty-seven species by name; Elton over two hundred. Given the differences in allowable length for each publication, 8,000 words for Bates and (ultimately) 50,000 for Elton, this works out to a slightly higher density of mentioned cases in the shorter paper. Working essentially from scratch over a period of a few months, Bates cited 46 sources and mentioned some earlier ones referred to therein. Two years later, Elton cited 297 sources, of which at least 33 (from 1956 and later) and up to 44 (adding those published in 1955) would not yet have been available to Bates. Both authors cited multiple-species, regional, or topical reviews about twice as often as monographs or other works dealing with individual taxa (usually species). As mentioned previously, their bibliographies included only eight sources in common, presumably representing the maximum number of “useful references” Elton found in “Man as an Agent.” Perhaps the most substantial of these was Henry N. Ridley’s The Dispersal of Plants. Viewed in such a limited statistical light, the two works seem roughly similar in many respects. But they were conceived very differently. Discussions of neobiota can be organized geographically, or by taxon, or by some ecological correspondence such as trophic level, or according to mode of transport, or by some measure of cultural (e.g., medical or economic) significance. All but the 449 Ibid., 801. 201 organizing criterion must then be recapitulated in every successive section. Bates chose the taxonomic option. His subheadings appeared in familiar “great chain of being” order: microorganisms, plants (deliberate and then accidental introductions), invertebrates, vertebrates (domestic[ated]; sport, food and fur; biological control; sentiment[al]). He closed with “the human habitat,” the meanings attributable to the phenomena discussed. Elton took another tack (described in detail in Chapter Four) maintaining and expanding on “Balance and Barrier’s” three-part format. His first (of nine) chapters, titled “The Invaders” was a tour-de-force litany of familiar and/or egregious case anecdotes. Like Bates, Elton closed with implications for humanity, but dwelled on different aspects: “The Reasons for Conservation” and “The Conservation of Variety.” Developments After 1958 By 1961, Marston Bates was recommending Elton’s book as “a lucid discussion” of neobiota.450 He credited Elton with defining the three-pronged justification for conservation: “the whole idea of trying to look at man's relations with nature in terms of ethics, esthetics and utility comes from Charles Elton.”451 In so saying, he retreated (or perhaps recoiled) from Elton’s “religion,” and reduced “practicality” to issues of human benefits, rather than Elton’s more expansive array of benefits and threats. Bates’s representation actually echoes 450 M. Bates, Man in Nature, (Englewood Cliffs, NJ: Prentice-Hall, 1961) 76n. 451 M. Bates, The Forest and the Sea (cit. n. 390), 268. 202 Aldo Leopold more strongly than Elton, but in the same (1960) note Bates overtly credited Leopold only with “the idea of an ecological conscience.”452 The Ecology of Invasions by Animals and Plants remained in print throughout Elton’s lifetime and was reissued in 2000 by the University of Chicago Press, thereby finally gaining an academic imprint.453 Elton’s book has so far been cited in at least 119 published scientific papers.454 In that form, Elton’s ecological authority has also buttressed many further technical and popular books, and a profusion of magazine and newspaper articles. Despite its more objective tone and formal symposium provenance, Bates’s “Man as an Agent in the Spread of Organisms” barely registers in the bibliography of today’s invasion biology for the simple reason that Bates did not describe the phenomenon as ‘invasion’, or as any sort of crisis. It has been cited only twelve times– an order of magnitude less frequently than The Ecology of Invasions– and four of the articles citing Bates also cited Elton.455 As shown in Chapter Two, Charles Elton’s attitude toward neobiota crystallized early in his career, and perhaps even before his career began. Elton’s growing scientific knowledge informed the way he expressed his concerns, but were not their basis. Anyone might be concerned by the relatively 452 I have seen no documentation or other evidence to suggest that Bates and Leopold ever met or corresponded. 453 Coincidentally, also the publishers of Man’s Role in Changing the Face of the Earth. 454 ISI Web of Knowledge Science Citation Index, queried 10 November 2006. 455 Ibid. 203 sudden appearance of large numbers of unfamiliar plants and animals; anyone might be alarmed further by reports that the phenomenon was becoming increasingly common. Marston Bates rarely saw an introduced plant or animal in the Colombian jungle. He approached the topic of neobiota as an ecological literature review. Lacking, or perhaps merely unburdened, by long personal involvement with the issue, or by the prospect of experiencing a real military invasion, he saw it in something approaching ecological terms. Marston Bates liked to point out that there was no profit to ecology in playing the game he called, “let’s pretend that man doesn’t exist.”456 But it seems that his own geographical isolation, combined with a relative disregard for convention, allowed him to overlook the power of religion and aesthetics in driving societal agendas, and even the science applied to them. Furthermore, he overlooked the power of nostalgia, and the fact that there is always a ready audience for a tale of a fall from grace, and a laying of blame for exile from the perfect garden. It seems that Elton knew better. Other Notable Syntheses Published between 1949 and 1959 “Man’s Role” was initially chosen as the topic of this chapter because of the unusual way Elton acknowledged it in The Ecology of Invasions. Subsequent research suggested that enough information existed to make comparing and contrasting the two works and their authors worthwhile. But Bates and Elton 456 See, e.g., Frank E. Egler "The Nature of Naturalization." Recent Advances in Botany. IX International Botanical Congress, 1959. D.L. Bailey, Ed. (Montreal: University of Toronto Press, 1961) Vol. 2, 1344; also the earlier appearing quotation from Bates’s 15 February 1955 letter to Elton. 204 were not alone in addressing these concepts during the 1950s. For example, K.A.G. Wodzicki’s 1950 Introduced Mammals of New Zealand was discussed in Chapter One. At least four more deserve mention in passing, as candidates for further investigation and expanded contextualization.457 In the late 1940s well-known naturalist and author Richard (“R.S.R.”) Fitter began collecting “an alarming array of facts”458 regarding introduced vertebrates in Britain, and in 1950 published an article titled “Man’s Additions to the British Fauna.”459 In 1959 he also released an expanded book on the topic, The Ark in Our Midst, which included the following acknowledgement: “I am … much indebted to an army of correspondents … whom I have pestered for information, or who have helped me in various other ways since Charles Elton first encouraged me to embark on this enterprise.” Listed again among the ranks of Fitter’s sixty-five member army was Elton himself. The extent of Elton’s encouragement and involvement in Fitter’s project is unclear, but it seems safe to say that as a man who openly valued rapid composition, he perhaps despaired of its completion after a year or two had elapsed. If Elton felt any duty to respect Fitter’s priority, it may have been diminished in the process. The two books are more complementary than 457 Other articles and books appeared between 1950 and 1960 making interesting but more restricted, taxon- or region-specific inventories and analyses of introduced biota, but there is insufficient space to discuss them all here. 458 R.S.R. Fitter, The Ark in Our Midst (London, Collins 1959), 7. 459 R.S.R. Fitter, “Man’s Additions to the British Fauna” Discovery 11 (1950), 58- 62. 205 competitive, and the fact that they ultimately appeared in fairly rapid succession suggests that Fitter was urged to completion when Elton took matters into his own hands. Fitter did not cite or acknowledge The Ecology of Invasions, nor did Elton cite Fitter’s article (or acknowledge Fitter). Herbert Levi of the University of Wisconsin wrote “Evaluation of Wildlife Importations” which appeared in the June 1952 issue of The Scientific Monthly.460 Levi’s major point was that even the best-laid introduction plans had rarely if ever been wholly successful. In the equivalent of about five full pages of text he briefly described approximately fifty cases of unintended consequences attending the transfer of animals, plants and disease organisms to new locations. Although his treatment was shorter than the others, Levi discussed hybridization and introgression more thoroughly. Levi cited early work by Elton (Animal Ecology, 1927) and Aldo Leopold (Game Management, 1933). Marston Bates cited Levi generally regarding the effects of intentional wildlife introduction, and also specifically for his discussions of crustacean introductions into European waters.461 Most, but not all of Levi’s cases later made appearances in Elton’s The Ecology of Invasions, but Elton did not cite Levi. Either Elton was unaware of the piece, or it did not meet the same unspecified threshold of recentness Elton attributed to Bates. 460 Herbert W. Levi “Evaluation of Wildlife Importations” The Scientific Monthly 74 (1952), 315-322. 461 Bates, “Man as an Agent,” (cit. n. 403) 797. 206 Carl H. Lindroth, Professor of Entomology at Lunds University, Sweden, published The Faunal Connections Between Europe and North America in 1957.462 He cited neither Bates nor Elton at any date, and neither the American nor the Englishman seems to have cited Lindroth afterwards. Although his second chapter (of three) was titled “The Human Transport of Animals Across the Northern Atlantic,” he also discussed and listed plant species. The book successfully restricted itself to trans-Atlantic transport but was in many ways more overtly grounded in ecological explanation, historical research and empirical natural history than the treatments by Bates and Elton. Lindroth’s account was far more detailed and data-laden than that in “Man as an Agent.” Although he referred at various points to “immigrants,” “synanthropes,” “inquilines,” “man’s constant followers,” “unwelcome passengers” and even “intruders,” Lindroth did not espouse a conservation philosophy, and so may have been doomed to relative obscurity. The final author I will mention was Canadian-born, University of Wisconsin geographer Andrew Hill Clark. He was responsible for two publications significant in this context. The Invasion of New Zealand by People Plants and Animals: The South Island, was based on his University of California (Berkeley) doctoral dissertation.463 “The Impact of Exotic Invasion on the Remaining New World Mid-Latitude Grasslands,” was prepared for and discussed in the same 462 Carl H. Lindroth, The Faunal Connections Between Europe and North America (New York: John Wiley and Sons, 1957). 463 Andrew Hill Clark. The Invasion of New Zealand by People Plants and Animals: The South Island, (New Brunswick: Rutgers University Press, 1949). 207 symposium session as Marston Bates’s “Man as an Agent.”464 Clark’s New Zealand book stuck to its regional focus, and his later paper offered a somewhat tortured rationalization for again dealing with two areas he already knew relatively well (grasslands in New Zealand and North America). But they both provided thoughtful accounts of the causes and effects of neobiota within their stated constraints. It is little more than a guess to suggest that Marston Bates reduced the potential redundancy of his symposium session’s offerings by avoiding specific reference to Clark’s work. It is more difficult to suggest that Charles Elton could have been unaware of Clark while simultaneously acknowledging Bates, since their papers appear in the same volume (and Clark cited The Invasion of New Zealand in his bibliography there). Like Elton, Clark favored the term “invasion,” which would seem likely to have attracted Elton’s attention. Several copies of Clark’s book occupy the stacks of Oxford libraries, but none occur in the Elton Library’s collection. By the 1950s, then, there were still several distinct motivations for studying neobiota. Charles Elton’s outlook was a personal conclusion, not an inevitable one. Marston Bates saw the phenomena as integral to studying human ecology. Carl Lindroth employed them as tools of archaeological analysis. Andrew Clark hoped his study would be “exemplary of the themes of 464 Andrew Hill Clark ,“The Impact of Exotic Invasion on the Remaining New World Mid-Latitude Grasslands,” in William Thomas, et al, Man’s Role in Changing the Face of the Earth, 737-762. 208 historical geography.”465 Richard Fitter wrote, to “a student of the problems of distribution and population, the opportunities afforded by the involuntary appearance of an animal in a new country are immense.”466 Herbert Levi warned that wildlife importations were rarely practical successes. The view from Oxford’s BAP was not the only view available, or necessarily the best one. Having looked at Elton’s precursors, his life and influences, and his competition, what remains is to more closely examine The Ecology of Invasions by Animals and Plants to determine which of these things left their marks on it, and illuminate the thoughts Elton actually pulled together in this book, whether for the first time or otherwise. 465 Clark, The Invasion of New Zealand, (cit. n. 463) v. 466 Fitter, The Ark in Our Midst (cit. n. 458), 13. Chapter 4: Dissecting The Ecology of Invasions by Animals and Plants Aside from the fact that the BBC had scratched his portentous title, “Ecology and the Millennium,” Charles Elton’s three radio lectures in the March 1957 were well received. Audiences found them to better than average examples of their kind. But they did not cause a stir. The term “infotainment” probably captures their essence, albeit anachronistically. Although Elton sometimes literally fled public attention, this outcome did not satisfy him. He decided to risk further notoriety by expanding the talks into a book, the first he had written primarily for a lay audience in twenty-five years. The previous one, Exploring the Animal World, had also followed hard on a series of radio lectures; so hard that it was little more than a transcript decorated with a few woodcuts. The new one was a more ambitious project. Documentation of their negotiations has not been found, but Methuen & Company of London, who published Elton’s tiny tome The Ecology of Animals in 1933, agreed to take it on. We do not know which party initiated the discussion. In his preface to The Ecology of Invasions by Animals and Plants (Invasions, following), Elton wrote “In this book I have tried to bring together ideas from three different streams of thought with which I have been closely concerned during the last thirty years or so. The first is faunal history… the second is [population] ecology… the third is conservation.”467 Taking his 467 Elton, The Ecology of Invasions (cit. n. 162), 13. 210 statement as a cue, this chapter summarizes Invasions, emphasizing those three matters, concentrating on structure, content and argumentation. Like a plot summary of Alice in Wonderland, the present chapter explains the structure, but only hints at the rhetorical power and appeal of Invasions. Elton’s intellectual style was critically important. To separate style and content in this case risks substantially misrepresenting and misunderstanding not only the way Elton wrote about his topics, but the way he actually conceived of them. One colleague fondly recalled, “He read widely outside science and had a dry sense of humour. I am grateful for his having introduced me, long ago, to the works of such as James Thurber and Damon Runyon.”468 Upon Elton’s death, two of his close acquaintances left us further hints: “the enormous breadth of his scholarship … never failed to impress his students,” and “his originality and vision were matched by [among other things] his gift for lateral thinking.”469 At least twenty times in Invasions, Elton invoked “non-science” sources, for example, Xenophon, Karel Čapek, Samuel Johnson, Charlie Chaplin, Shakespeare, Isaiah, and someone identified only as “a great literary critic,” (but in fact, his father Oliver). Only a few can be mentioned in context, below; but such writers and their works were seemingly ever-present in Elton’s mind. Taking another cue then, this time from Aldo Leopold (whose textbook Game 468 Harry Thompson, 1991, quoted in Dennis Chitty, Do Lemmings Commit Suicide? (cit. n. 187), xvii. 469 Kitty Paviour-Smith, “Obituary: Charles Elton” The Independent (London) 13 May 1991; Max Nicholson, “Rebellion Against the Ratpack: Obituary of Charles Elton, FRS” The Guardian (London) 13 May 1991. Both accessed online at web.lexis-nexis.com, 4 September 2006. 211 Management recommended that users set it aside and read Elton’s Animal Ecology first): readers of this treatment would benefit by reading (or re-reading) Invasions before proceeding.470 The book is brief, engaging and readily available. Invasions was published on 10 July 1958. With about 100 octavo pages of text, it was the fifth and last of Charles Elton’s “little” books. Three of the other four (Animal Ecology, The Ecology of Animals, Animal Ecology and Evolution) were intended as textbooks, but they were published between 1927 and 1933, when animal ecology could muster few specialists. By 1958, ecology had significantly expanded, exhibiting both methodological subdisciplines and philosophical factions. Elton reserved his final “big” book (The Pattern of Animal Communities) for ecologists; but with Invasions he turned outward, attempting to extend or capitalize on the rare star turn as a public intellectual begun with his “Balance and Barrier” radio lectures. Only one advertisement for the book was found (fig 4.1), in the British Ecological Society’s The Journal of Ecology. Its tone is distinctly Eltonian, but perilously pedantic as an appeal to colleagues who knew perfectly well who Elton was. By almost any standard, this was exactly the wrong place for such an advertisement to appear. But aside from the broadcast lectures that preceded it, and possibly word of mouth, the advertisement was the first impression of the book provided to potential readers; so it appears first here, as well. 470 See Chapter Two 212 Figure 4.1 Advertisement for The Ecology of Invasions in unnumbered back pages of The Journal of Animal Ecology for July 1958. An exhaustive search was not attempted, but after some effort I found no evidence of Invasions being actively marketed to the intended audience. It apparently received no attention in either the British or American general interest magazines that typically carried such content. No reviews or articles regarding Invasions were recorded in book review indexes or The Readers Guide to Periodical Literature. A small, bibliography-style “New Books” notice appeared, along with many others, in the November 1958 Journal of Animal Ecology. Another was published in the 7 November 1958 Science, showing that Wiley was publishing Invasions in the United States. 213 Several reviews of the first edition soon appeared in scientific and professional journals. These and less comprehensive (but contemporary) comments are discussed later in this chapter. Organization Structurally, Invasions was more a semi-integrated trilogy rather than a single elaborate argument. The book’s organization followed Elton’s “three streams of thought,” and was probably intensified by its genesis as a radio series composed in three potentially stand-alone episodes. Making the transition from broadcast to book, Elton mostly added cases and examples rather than substantially reorganizing and integrating the radio scripts. Elton’s first five chapters comprised “book one” of the Invasions trilogy. There he concentrated on faunal history, a term Elton preferred to “zoogeography” which he scorned as “an enormous and indigestible subject.”471 Chapters one and two reworked and expanded Elton’s obscure 1943 paper “The Changing Realms of Animal Life.”472 They introduced the problems that concerned Elton, and provided some paleontological background information. Chapters three through five consisted largely of geographically sorted case histories. They dealt, respectively, with the recent faunal histories of continents, remote islands, and oceans, and to some extent compared and contrasted the issues Elton believed were significant or unique to each of theses categories. 471 Elton, Invasions (cit. n. 162), 35 472 See Chapter Two. 214 Chapters six and seven of Invasions comprised “book two,” introducing a few basic concepts from ecology, and adding further cases and anecdotes exemplifying these concepts. After defining the problem with books one and two, he set out to solve it in book three. He also made explicit the philosophies and values tacitly underlying his many case histories. Invasions chapters eight and nine thus completed the trilogy by explaining Elton’s arguments for conservation action, and then for the specific sorts of actions Elton felt were needed in Britain. Chapter eight included only a few new examples, and revisited some already introduced, but Chapter nine added none at all. Book One, Part One: A Very Explosive World The explicit thesis of the first two chapters, and “one of the first themes of [the] book” was: “if we are to understand what is likely to happen to ecological balance in the world, we need to examine the past as well as the future.”473 However, Invasions opened with a partial thesis that can only be termed blatantly alarmist. “Nowadays we live in a very explosive world,” it began, “and while we may not know where or when the next outburst will be, we might hope to find ways of stopping it or at any rate damping down its force. It is not just nuclear 473 Elton, Invasions (cit. n. 162), 33. 215 bombs and wars that threaten us … there are other sorts of explosions, and this book is about ecological explosions.”474 After mentioning a few dread epidemics and agricultural disasters (e.g., influenza, bubonic plague, malaria; the Irish potato famine) Elton posited that such events were more common than formerly, and that “we need to understand what is causing them and try to arrive at some general viewpoint about the whole business.”475 He followed with a taxonomic smorgasbord of seven brief “case histories of species which were brought from one country and exploded into another.”476 These included an insect, a fungus, a bird, a mammal, a plant, a fish, and a crab. “The real thing,” Elton wrote, “is that we are living in a period of the world’s history when the mingling of thousands of kinds of organisms from different parts of the world is setting up terrific dislocations in nature.”477 Chapter one ended on a note that exemplified the catholic eclecticism of Elton’s thinking. “We might say, with Professor Challenger, standing on Conan Doyle’s [1912] ‘Lost World’ with his black beard jutting out: ‘We have been privileged to be present at one of the typical decisive battles of history — the battles which have determined the fate of the world.’” Then playing the line out 474 Ibid., 15. Britain detonated its first atomic bomb in the Pacific about six weeks after Elton’s final broadcast, and went on to test a fusion bomb later the same year, a month after the orbital debut of Sputnik 1. 475 Ibid,. 18. 476 Ibid,. 19-20. 477 Ibid,. 18. 216 further, “But how will it be decisive? Will it be a Lost World? These are questions that ecologists ought to try to answer.”478 Elton began chapter two with a brief counterfactual, describing his conception of a world “where there were no barriers to [the geographical] spread [of animals].” He then pointed out that “quite a large number of species are able to achieve a world-wide distribution as it is, either because the ecological barriers that hold in others are not barriers to them,” giving several examples.479 But the remainder of the chapter combined descriptive paleontology and biogeography to support the idea that “a great many other plants and animals never had the opportunity of ranging over the whole world … these became cooped up, as it were, in various regions for long enough to change profoundly and leave a permanent mark on the composition of flora and fauna.” Elton recognized that he was oversimplifying faunal history and immediately qualified the foregoing statement by describing two more “processes 478 Ibid,. 31-32. Elton’s quotation (from Doyle’s Chapter 14) included a tacit ellipsis and significant decontextualizing. The battle Professor Challenger witnessed pitted “ape men” against “Indians:” Challenger's eyes were shining with the lust of slaughter. "We have been privileged," he cried, strutting about like a gamecock, "to be present at one of the typical decisive battles of history — the battles which have determined the fate of the world. What, my friends, is the conquest of one nation by another? It is meaningless. Each produces the same result. But those fierce fights, when in the dawn of the ages the cave-dwellers held their own against the tiger folk, or the elephants first found that they had a master, those were the real conquests — the victories that count. By this strange turn of fate we have seen and helped to decide even such a contest. Now upon this plateau the future must ever be for man." 479 Elton, Invasions (cit. n. 162), 33-34. 217 that increase the complexity of the distribution pattern as we see it now.” These were the contraction of some formerly large ranges and apparent, prehistoric “remingling of faunas” after “long periods of isolation.”480 He accounted for these by invoking geographic dynamism (e.g., changes in sea level and continental juxtaposition) without speculating as to its causes.481 Although Elton was aware of Alfred Wegener’s 1912 continental drift hypothesis and had even lectured on it, that idea would not begin its rehabilitation and refinement by geophysicists into plate tectonic theory until 1961-1962.482 Lacking this essential, central dogma, his paleontological arguments in chapter two cohered poorly. Alongside his problematic prehistory, Elton interposed a celebration of Alfred Russell Wallace and “his wonderfully rich experience of some of the most exciting facts of zoogeography.”483 Elton described Wallace’s six faunal regions, and added another of his own devising, to cover “the vast Eastern Pacific beyond the Australian continental arc, with its Milky Way of islands.” One can imagine Elton whispering, with a nod and a wink, as he confided to his readers, “There 480 Ibid,. 34. 481 In one noteworthy instance, (Ibid,. 36) Elton called several vaguely Cretaceous or post-Cretaceous events that apparently isolated various continents “a particularly important accident of history,” seemingly heedless of the implications that statement might have for any subsequent normative claims. 482 Crowcroft, Elton’s Ecologists (cit. n. 184),140; Anonymous, “Plate Tectonics: The Rocky History of an Idea” (University of California, Berkeley, Museum of Paleontology, 1997) accessed online (31 August 2006) at http://www.ucmp.berkeley.edu/geology/techist.html. 483 Elton, Invasions (cit. n. 162), 36; See Chapter One. 218 are, in fact, seven great ‘realms’ of life.”484 Elton’s discussion of Wallace seems very deferential, giving a distinct impression of the ‘just-so’ story. To him, Wallace’s Realms were not only received truth, but received propriety. An earlier, related passage carried the same message. “When one was a child, [the distinct faunas of different realms were] very simply summed up in books about animals. The tiger lives in India. The wallaby lives in Australia. The hippopotamus lives in Africa.”485 Here, Elton’s very “Atlantic” nostalgia for childhood security intertwined itself with his received antiquarian fancy of an uncomplicated, “pre-Atlantic” world.486 This section was awkward in other respects. Charles Darwin received a brief but honorable mention, as Elton related how Darwin discovered fossils of extinct mammals representing several orders and families in southern South America. These anecdotes are perfectly at home in chapter two, but Elton’s familiarity with them raises a question about his complete omission of Darwin’s almost concurrent accounts of finding European plants growing abundantly in the same region.487 The wry quips and culturally familiar analogies that characterized Elton’s other chapters were noticeably sparse in chapter two. In fourteen pages, references to a “chinchilla the size of a rhinoceros,” and to faunal 484 Ibid,. 37. 485 Ibid,. 30. 486 In a post- Balance and Barrier letter to John Simons, Elton declined a BBC request to speak on the literally pre-Atlantic subject of continental drift, protesting “I do not know enough and am no good at public discussion.” (C.S. Elton to J. R. Simons, 4 July 1958; BBC:ETF). 487 See Chapter One. 219 history “emerging like a photograph in a slow developer” were all he could manage.488 Finally, Elton neither attempted a summary conclusion, nor set the stage for chapter three. Combined with difficult topics and theoretical weaknesses, this made chapter two relatively inaccessible. It is easy to imagine a general reader (or an ecologist) skimming a bit here and there Book One, Part Two: Continent, Island, Ocean The next section commenced with a brief prolegomenon. This complained of the way newly established species complicated “ecosystems already difficult enough to understand, let alone control,” appropriated Edward Gibbons’ accusation that his materials, and not merely his methods, were deficient, and then lauded the United States Department of Agriculture (USDA), “the first to bring some sort of method and order into this field.”489 Then Elton disclosed his concerns, but muddied them with a complicated and arcane analogy. “No one really knows how many species have been spreading from their natural homes, but it must be tens of thousands … if we look far enough ahead, the eventual state of the biological world will become not more complex but simpler — and poorer. Instead of six continental realms of life, there will be only one world.” He envisioned this “one world” as a collection of tanks, each filled with different, complex chemical mixtures. Human intervention was rapidly intermingling their contents, during which process he presumed some compounds would “disappear” through recombination. He saw this as a net loss. Breaking the 488 Elton, Invasions (cit. n. 162), 39, 41. 489 Ibid,. 50. 220 frame, he wrote “although there is a Law of the Conservation of Matter, there is no Law of the Conservation of Species.”490 For the equivalent of twelve pages, or about one-eighth of the total text of Invasions, Elton discussed insect pests; mostly in North America, and mostly relying on reports from the USDA, whose expertise he touted previously. All were regionally, economically important cases. Elton felt it necessary to deny that he chose them as such, but rather, because they “illustrate the ideas of this book, or … happen to have good maps of the invasions of continents by foreign species.”491 About halfway through the insect list, Elton gave his readers a break, and proposed another analogy: “The disadvantage of describing invasion only by examples, however famous, is that this does not quite convey the tumult and pressure of species that are escaping from the confinement of their ancestral continents to range the world.” After quoting lines eight through twelve of Walt Whitman’s poem of gathering waters, “As Consequent, Etc.,” Elton explained: “Whenever we know the history it starts with a very small nucleus of population, growing to an ‘Autumn Rivulet’ and then not infrequently to a flood. And when the population has got that far, its movement is seldom absolutely checked except by natural limits of the environment.”492 Then he returned to cases. 490 Ibid,. 51. 491 Ibid,. 50. 492 Ibid,. 61. 221 The last two “Continents” pages dealt with vertebrates. He contrasted these with “insect immigrants” (an uncharacteristically mild epithet) by pointing out that the insects were “introduced by mistake,” but the vertebrates (barring rats and mice) were “brought intentionally in the first instance, though many of them have become extremely harmful or unpopular afterwards.”493 He declared his chapter one discussion of muskrats to be a representative case, and then provided an unadorned list of several more animals and the places they were newly inhabiting and affecting, finishing with the release of tropical aquarium fishes into the Florida Everglades. Like the previous chapter, this one ended abruptly. “The Fate of Remote Islands,” opened with a quote from Captain James Cook, on arriving at Easter Island in 1774: “‘Nature has been exceedingly sparing of her favours to this spot.’” Elton concurred in a fashion that raises questions rather than answering them: “This was exactly true, for Nature had only with great difficulty managed to get there at all.”494 The balance of the chapter compared the former, present, and sometimes projected future biotas of various island groups. Case histories of species carried to Tristan da Cunha (37°S, 12°W), Hawaii, and New Zealand accounted for bulk of this information; perhaps not least because accounts of them were published and readily available. Although Elton had never visited any of the islands mentioned in the chapter, 493 Ibid,. 73. 494 Ibid,. 77. 222 their “fate” in this regard seemed to arouse his indignation more than any other single aspect of his topic. After briefly describing pre-Cook Hawaii, Elton queried himself, “what has been the fate of this marvellous flora and fauna?” and expended six more pages in answer. His expression became more emphatic. One “voracious,” “ubiquitous,” “scourge of native insect life,” “alone has accounted for untold slaughter.”495 “There can be few invading species which become such a menace to motor traffic that they cause cars to skid on the roads!”496 “It is quite an exchange and bazaar for species, a scrambling together of forms from the continents and islands of the world, a very rapid and efficient breaking down of Wallace’s Realms and Wallace’s Island Life!”497 Anticipating Paul and Anne Ehrlich’s 1981 aeronautical rivet-popping analogy: “It mitigates the fate of remote islands in this century if some of their species are saved from the wreckage, like the few survivors that clamber out of a smashed aircraft.”498 Elton spiced his New Zealand case histories with another vehement contention. “No place in the world has received for such a long time such a steady stream of aggressive invaders.”499 495 Ibid,. 82. 496 Ibid,. 85. 497 Ibid,. 83. 498 Ibid,. 88-89. 499 Ibid,. 89. 223 Before closing, Elton framed a litany of questions that perfectly anticipated Robert MacArthur and Edward O. Wilson’s queries regarding “island biogeography,” but were (problematically, in my view) unacknowledged by the American duo.500 “What is the full ecosystem on a place like Guam or Kauai or Easter Island? How many species can get along together in one place? What is the nature of the balance amongst them?”501 Elton bade farewell to the islands with a peculiar anthropological digression, quoting at uncommon length from both primary and secondary sources. I would like, however, to leave [this] subject with a back-glance at a more pleasant and balanced ecological world, before Atlantic civilized man crashed into this remote galaxy of island communities. In that age, when numbers of human beings were regulated by customs, often harsh enough, but meeting the end desired, a great many of the Pacific Islands were inhabited by quite large numbers of a small species of rat, derived from a Malayan form, and evidently brought by the Polynesians in their great migrations eastwards and southwards some hundreds of years ago. Rattus exulans. … is a small rat, much gentler in habits than [black and brown] rats.”502 He continued for a further page, extolling the significance of these kinder, gentler rats for island societies: On the island of Raratonga, in Mid-Pacific, they were highly prized for sport. ‘In those days—ere the cat had been introduced—rats were very plentiful. Rat hunting was the grave employment of bearded men, the flesh being regarded as delicious.’ And on the Tonga Islands about 1806, the King and court used to go out and shoot the rats along the forest 500 MacArthur and Wilson developed “island biogeography” theory between 1962 and 1967. 501 Elton, Invasions (cit. n. 162), 89. 502 Ibid,. 92. 224 paths, using huge bows and arrows six feet long: ‘Whichever party kills ten rats first, wins the game. If there be plenty of rats, they generally play three or four games.’ There were elaborate rules, as we should now have for football or hunting: precedence, offside, and—a wonderful dispensation—if you shot a bird you could count it as a rat!503 Having introducing the “pleasant and balanced ecological world” when humans still died off at prudent rates, he finally closed, “we will hope that … research will find out how the broken balance can be restored and protected.”504 “Changes in the Sea” was the only chapter of Invasions adorned with an epigraph. Elton quoted six lines of the epic “fragment” Hyperion (1820) by John Keats, omitting the four italicized below. It seems to have been selected mainly for its vocabulary, rather than its relevance, given the need for selective omission. For though I scorn Oceanus’s lore, Much pain have I for more than loss of realms: The days of peace and slumberous calm are fled; Those days, all innocent of scathing war, When all the fair Existences of heaven Came open-eyed to guess what we would speak: — That was before our brows were taught to frown, Before our lips knew else but solemn sounds; That was before we knew the winged thing, Victory, might be lost, or might be won. Although Elton had extensively studied pond life as a child and then a University student, marine biology could not be called one of his strengths. Nevertheless, he plunged in, identifying three kinds of pertinent “human actions … First the digging of new canals. Secondly, accidental transport on ships. And 503 Ibid,. 93. 504 Ibid, 225 thirdly, deliberate introductions.”505 These cases were stated matter-of-factly. Elton exhibited little personal sympathy for marine life of the sort he had evidenced for island biotas. There were (and are) only two “new” canals connecting marine biotas, in Panama and Suez. Acknowledging that each included lakes, one freshwater (Gatun, in Panama) and the other hypersaline (The Great Bitter Lake), Elton nevertheless distinguished between chemical physical barriers to migration. He identified only one species that had completely crossed Panama, but wrote of “a fairly strong contingent of animals [that] has managed to pass from the Red Sea to the Gulf of Suez and spread into the Mediterranean.”506 Elton mentioned several hull-fouling, coastal organisms that apparently crossed oceans attached to ships, and speculated about the possibility of carriage in ballast-water tanks without giving any examples. But he devoted the bulk of this discussion to intentionally transported and introduced species. He saw farming of oysters and clams, including the nearly cosmopolitan exchange of breeding stock between Asian, American and European growers, as tantamount to domestication; oysters were “a kind of sessile sheep, that are moved from pasture to pasture in the sea.”507 Like other domestic animals, their transportation involved inadvertent movement of associated species, leading to a 505 Ibid,. 94. 506 Ibid,. 94-95. 507 Ibid,. 100. 226 digression on that point. Elton mentioned a jumble of these accidental cases, finally returning to intentional introductions of game fish in several oceans. As a middle-class Englishman, Elton may have associated angling with upper-class indulgences, but whatever the reason, after providing statistics on the effort expended on capturing one transplanted game species, he closed with a pot-shot at sportsmen: “A world that begins to assess its recreation in manhours probably cares fairly little about the breakdown of Wallace’s realms.” Then, relating an aquatic version of his questions about island populations, he closed the discussion of faunal history and turned to ecology. Book Two: Popularizing Populations, and a Feeling of Feeding Elton began “book two” by summarizing his thesis for “book one,” first in military parlance, and again, allegorically, with an extended quote from H.G. Wells’ Food of the Gods. He then added the new partial thesis that most “invasions … never happen” because “they meet with resistance” by “man or by nature or by man mobilizing nature in his support.” But even though chapter six was titled “The Balance Between Populations,” Elton mustered several more pages of martial imagery before really addressing ecology. The ecological nub of Invasions boiled down to asserting that competition occurs between populations, sometimes resulting in one population occupying an area where another had formerly been found. But Elton did not go so far as to define “population.” And he was extremely cautious, even equivocal about claiming that direct or “interference” competition between populations could 227 actually be demonstrated. “I have set out … examples where competition is for space to live and eat or breed, because they are the simplest ones to understand. But it is likely that competition is usually a far more complicated matter.” Even further, he noted that “replacement of one species, or part of the populations of one species by another” may follow from “a whole string of causes and effects that are very hard to trace.”508 Only in passing here did Elton mention an “ecological system” (not an “ecosystem”) and “communities,” never substantially attempting to relate them to species or populations. Even though he emphasized the existence of ecological barriers between populations, and the significance of changing their effectiveness, Elton saw no real prospect of repairing them, of erecting surrogates, or of finding “new technological means of suppressing this plant or that animal.”509 [W]e have to accept the proposition that invasions of animals and plants and their parasites—and our parasites—will continue as far as the next Millennium and probably for thousands of years beyond it. Every year will see some new development in this situation. That is a way of saying that the balance between species is going to keep changing in every country. Quarantine and the massive campaigns of eradication are ways of buying time—though they are valuable and necessary, they are also extremely expensive. It takes so few individuals to establish a population, and such a lot of work to eradicate them later on. [italics and capitalization Elton’s]510 This comprehension (or resignation) is one of the keys to understanding the philosophy he espoused later in “book three.” Elton could see the moving finger 508 Ibid,. 122. 509 Ibid,. 110. 510 Ibid,. 112. By 2006 Elton’s estimate appeared to be conservative. 228 of nascent economic globalization writing the foreseeable future of life.511 He didn’t like what he saw, and hoped to “damp it down a bit,” but understood that neither piety, nor wit, nor tears would call it back. Even though Elton was thus apparently resigned to ongoing ecological change for the foreseeable future, a nostalgic sense of “Paradise Lost” pervades Invasions.512 Modern nostalgia has been described as “mourning for the impossibility of mythical return, for the loss of an enchanted world with clear borders and values;” and “longing for that shrinking space of experience that no longer fits the new horizon of expectations.”513 Some commentators differentiate nostalgia from “antiquarianism;” the former composed of personal memories and the latter a yearning for an unexperienced but appealing historical setting.514 In either case, memory being both idiosyncratic and fallible, the past is inevitably reconstructed from its most attractive aspects. Acknowledging the (re)constructed aspects of nostalgia is important here because Elton appealed to both a pre-Edwardian conception of the ruling class as sportsmen-hunters, and a pre-technologically “balanced” nature. These constituted prime examples of nostalgic construction, hearkening “back” to (1) an 511 According to the online edition of the Oxford English Dictionary, the term “globalisation” first appeared in print in 1959. 512 Svetlana Boym, The Future of Nostalgia (New York: Basic Books, 2001), 3. 513 Ibid., 8; 10. Nostalgia was originally a medical diagnosis. Swiss physician Johannes Hofer coined the term in 1688; among its first diagnosed sufferers were “various displaced people of the seventeenth century” including soldiers deployed abroad for extended periods. 514 See, e.g., Fred Davis, Yearning for Yesterday: A Sociology of Nostalgia (New York: The Free Press) 1979. 229 experience he did not actually share, either as an Englishman or as a member of the Raratongan culture he described in “book one,” and (2) by repeatedly invoking nature’s “balance” a concept and condition he had long before repudiated (see Chapter Two) as a naive fiction, never to be experienced by anyone. Did this signify a relapse into pre-ecological thinking? This seeming retrogression to extolling a “balanced” nature needs further investigation and analysis; more than can be undertaken here. The various comments he made regarding balance in Invasions seem inconsistent at best. Some describe an essentially static condition. Some of them reified the concept of a natural balance that always, eventually sorts itself out. Others suggested that balance is simply a desirable condition human managers should impose on nature, or at least encourage from it. And one saw him metamorphose from strident Cassandra to reflective empiricist in the space between two sentences: “When new species arrive and spread, even if they do not have the appearance of the explosive invader, they may herald the onset of future changes in the balance between populations. The complete unravelling of any of these relationships will be an interesting but often very difficult task.”515 Elton’s “populations” chapter also included a quirky digression regarding failed quarantine efforts. He gave two “close to home” examples. The first was an unidentified personal friend whose Egyptian souvenir shirt had palm nut 515 Elton, Invasions (cit. n. 162), 124. 230 buttons harboring live beetles. The second was Elton’s own experience, indexed in Invasions under “acorns, a cautionary tale.”516 It has a familiar and yet hollow ring: I learned how easy it is to bring in a foreign insect when I carried home a few large American acorns from Wisconsin just before the War. I only wanted to have them on my desk for mementoes. A few days after I got back some chafer beetle grubs emerged from the acorns. Of course I dropped the whole lot into boiling water to kill them instantly, and that was the end of it. When the Customs officer had asked me whether I had anything to declare, it never occurred to me to say ‘acorns,’ and I am not sure that he would have been interested if I had.”517 These were undoubtedly the bur oak acorns Elton expropriated during his 1938 visit to Aldo Leopold’s “Shack.”518 But did he carry them home to admire, as stated here, or to propagate, as noted at the time? Did he collect enough to serve both purposes? Bur oak acorns are aesthetically interesting (Fig 4.2), so saving a few as desktop knick-knacks was a plausible motive. But in a parable about carelessness, in a book decrying introductions, Elton seems to have fraudulently disavowed his original motive: to plant American oak trees in England.519 Furthermore, asserting that he immediately referred the wormy nuts to a steaming auto-da-fe is also suspect. Elton’s curiosity regarding the emerging 516 Ibid,. 175. 517 Ibid,. 111. 518 Cf. Chapter Two. 519 In 1938, Elton was “Garden Master” for Oxford’s Corpus Christi College; he was also responsible for the grounds shared by his home at 61 Park Town and adjacent addresses. Inquiries have been initiated to determine whether there are extant bur oaks at either location, without result at the time of this writing. 231 insects would likely have prompted him to pick up a magnifying glass and a reference book. He certainly knew that the potentially problematic larvae could be utterly contained by simply “drop[ping] the whole lot” into a jar and screwing on the lid. This lapse recalls an earlier episode, Elton’s wartime memo to Arthur Tansley, in which he excoriated the behavior of an ecologist who trapped a population of island mice to the brink of extinction, admitting only later, privately (coincidentally, to Leopold) that Elton himself did this deed.520 We can only speculate on the extent to which such chinks in his “armor of truth” admit any additional light into Elton’s motivations and practices. Perhaps after twenty years Elton had forgotten his original intentions, and the existence of his notes. But if we merely credit him with accurate recall of a personally significant event, then he had three alternatives. He could leave his cautionary tale untold. He could trust his audience, tell the original story and apologetically take his knocks for temporary absent-mindedness. Or he could dissemble, buttressing his apparent moral authority and the associated credibility of his general claims. Whatever the decision process, he chose the latter. The story stood unchallenged for fifty years, and will likely be dismissed by Elton’s apologists with a shrug. But because he, himself intertwined and invested it with his personal scientific and moral authority, it casts a pall that could extend over the entire body of his scientific and conservation work. 520 See Chapter Two. 232 Figure 4.2. Bur oak acorns, shown approximately natural size.521 David Cox, author of a 1979 dissertation on Elton and the Oxford Bureau of Animal Population, recently suggested to me that Elton might have been jealous of Aldo Leopold’s exalted place in the pantheon of environmentalist authors.522 As shown previously, their relationship was already growing more 521 Photo adapted from Anonymous, Texas Tech University class materials accessed online at www.pssc.ttu.edu/pss1411cd/PLANTID/buroak/buroak.htm. 13 September 2006. 522 David Cox, personal communication, 16 August 2006. 233 distant by 1948. This might help explain (at least) why Elton omitted Leopold from the story instead of celebrating their relationship. On the other hand, perhaps he merely omitted Leopold out of discretion. The American was long dead, and there was no blatant privacy issue, but Elton had a peculiar penchant for secretiveness, making every personal anecdote difficult to verify. “New Food Chains for Old” began with a strong statement. “The natural living world is arranged in very complex channels of supply that are known as food-chains.”523 In a rare acknowledgement of another ecologist’s conception, Elton described this “cycle of supply” by quoting Alfred Lotka’s 1925 view: “one great world-wide [energy] transformer … one vast unit, one great empire.” But he even more prominently promoted an older conception, a 1556 drawing by Pieter Bruegel the Elder (Fig. 4.3). Its proverbial Latin title amounts, roughly, to “Big fish eat little fish.” As Elton explained it, “altogether this picture tells one more and makes one feel more about the supply lines of nature than any amount of formal logic might do.”524 After providing this “nightmarish insight,” Elton quickly turned to the chapter’s real thesis: “[W]ith land in cultivation, whether pastoral, ploughed, or gardened, the earnest desire of man has been to shorten food-chains, reduce their number, and substitute new ones for old.”525 He provided another litany of 523 Elton, Invasions (cit. n. 162), 125. 524 Ibid,, 126-127. 525 Ibid,. 127. 234 examples, listing who was eating whom, and describing how some such relationships were changing. Figure 4.3. Grandibus exigui sunt pisces piscibus esca reproduced poorly in early editions of Invasions (and worse in the 2000 facsimile) but clearly evidenced the idiosyncratic “foxing” stains on the original Bruegel drawing, part of the Albertina Graphic Collection, Vienna. But Elton deviated from the established pattern by discussing the advent of chemical herbicides and pesticides, specifically calling out “the new dusts and sprays” of DDT, parathion and 2,4D. Dripping with sarcasm, he wrote, “With this equipment … the applied biologist seeks to bypass all the irritating and complex interactions of natural populations, in fact simply sweep away natural food-chains altogether, leaving the crop plants to give an ordered and useful appearance to the landscape.” His discussion focused on two drawbacks of these chemicals: 235 early signs that target species were developing resistance, and what would later be called “non-target” or unintended effects on other species. He ultimately reduced his view of this chemical onslaught to a memorable metaphor: an “astonishing rain of death.”526 Book Three: Why We [Should] Fight Book three reveals Elton in the process of developing his “general viewpoint,” refining it, and convincing himself of its appropriateness. Instead of case histories it was dominated by a severally reiterated rationalization of Elton’s concerns and calls to action, based on a three-legged system of values, the conceptual “tripod” of ethical, aesthetic and economic conservation criteria he had formulated with Aldo Leopold.527 In the decade following Leopold’s death, Elton further developed and refined the tripod to suit his personal outlook, but not to the extent of revolutionizing it. He ostensibly labeled the first leg of his own tripod “religion,” rather than ethics. But there is no reason to suppose that he was allied with any religious orthodoxy, or a well-developed but idiosyncratic religious belief. He quoted once from the Old Testament (Isaiah 5:8), following immediately with a mildly parodic allusion to Matthew 4:4 (“man cannot live on gloom alone”). Either verse could plausibly arise in an educated English conversation. The Matthew quote has long been idiomatic. The first, though less familiar, was part of a standard, Church of England, Morning Prayer lesson and thus potentially familiar to many 526 Ibid,. 138-142 527 Cf. Chapter Two. 236 of his intended readers. The words served him well, but as was often the case, Elton fairly ripped them out of context: “‘Woe unto them that join house to house, that lay field to field, till there be no place, that they may be placed alone in the midst of the earth!’” 528 Elton’s illustration of religious duty, the “quite dangerous” practice of grain warehouse managers in India providing drinking water for rats, does not suggest that he held religion in high regard. He also quoted Albert Schweitzer regarding the problem of limiting ethical regard to humans, sans reference to deity or transcendence.529 So it is difficult to say exactly why Elton chose to invoke religion, unless he felt it to be a more palatable conceptual meal for the British public than interspecific ethics. Like Leopold, Elton called his tripod’s second leg “aesthetic” (though the Oxford man opted for the opening diphthong, rather than the rude midwestern “e”). More substantially, Elton’s expanded the concept to “aesthetic and intellectual.” He explained this in terms that mainly applied to his immediate family: “You can say that nature — wild life of all kinds and its surroundings — is interesting, and usually exciting and beautiful as well. It is a source of experience for poets [his wife Joy Scovell] and artists [his mother Letitia, his brother Leonard], of materials and pleasure for the naturalist and scientist [Charles himself and his late brother Geoffrey].” To all of which he appended a terse sentence fragment: “And of recreation.” Given Elton’s previously evidenced 528 Elton, Invasions (cit. n. 162), 143. 529 Ibid,. 144. 237 disdain for mere recreation, and we can imagine this phrase emanating from behind a grudging scowl. “In all this the interest of human beings is decidedly put first,” Elton wrote, effectively collapsing any distinction that could have been made between individual and collective motivations.530 The third and sturdiest leg of Elton’s tripod collected “practical” considerations, expressed as a neoMalthusian concern about human kind exceeding planetary limits. Since Thomas Robert Malthus’ infamous 1798 postulate that food supplies could only grow arithmetically while human populations grew geometrically, various authors have warned of dire, more or less imminent resource shortages. Elton, in turn wrote that the practical question of “land, crops, forests, water, sea fisheries, disease, and the like … seems to hang over the whole world so threateningly as rather to take the light out of [religious, aesthetic and intellectual arguments for nature conservation]. The reason behind this, the worm in the heart of the rose, is quite simply the human population problem. The human race has been increasing like voles or giant snails, and we have been introducing too many of ourselves into the wrong places.”531 Here, Elton also applied an unusual rhetorical strategy, appealing to six “hair-raising,” “fairly recent” books, not as substantial authorities for his 530 531 Ibid. Ibid. “The worm in the heart of the rose” was an unacknowledged quotation from the twentieth-century epic poem Savitri by Sri Aurobindo, based on the Mahabharata. 238 arguments, but merely juxtaposing their titles as a sort of moody free verse. 532 He left all but their authors unidentified and their specific contentions unstated, assuring us only that these were “serious works— not written by cranks: Road to Survival, The Rape of the Earth, Our Plundered Planet, The Geography of Hunger, Resources and the American Dream, The Limits of the Earth.”533 He added one more, this time explaining whose, and why. “Also The Estate of Man, in which Michael Roberts suggested that we are reaching the limit of the supplies of inherited talents needed to cope with these problems.”534 All but one of the listed books are neoMalthusian jeremiads. Copies of six remain in the Elton Library at Oxford, which constitutes the material detritus of the BAP.535 Josue de Castro’s The Geography of Hunger broke the mold; Elton would have found little of it quotable for his purposes. But he might easily have ghost written its jaded opinion of modern ecological practice: “Such a scientist as [Jean Henri] Fabre, who could lie on the ground for whole days at a time to watch the 532 It might be pointed out that “hair-raising” was a purely nostalgic idiom for Elton by 1957; coincidentally, according to the Oxford English Dictionary, the phrase came into use about the time Elton was born. 533 Elton, Invasions (cit. n. 162), 144. In order of appearance there: William Vogt, Road to Survival (New York: William Sloane Associates, 1948); G.V. Jacks and R.O. Whyte, The Rape of the Earth (London: Faber and Faber,1939); Fairfield Osborn, Our Plundered Planet (Boston: Little, Brown, 1948); Josué de Castro, The Geography of Hunger (Boston: Little, Brown, 1952); Samuel H. Ordway, Resources and the American Dream (New York: Ronald Press, 1954); and Fairfield Osborn, The Limits of the Earth (Boston: Little, Brown, 1953). 534 Elton, Invasions (cit. n. 162), 144; Michael Roberts, The Estate of Man (London: Faber and Faber, 1951). 535 Oxford University Libraries OLIS Catalogue, queried online 18 July 2006. 239 ways of ants and other insects, is a real rarity in modern life. The naturalist of today is little interested in how living things live — more in their behavior under complicated laboratory tests and their appearance, killed and quartered, under his powerful microscope. Yet it is certain that renewed exploration in the immense preserve of nature, carried out with love and patience, would reward us with new food resources whose existence we do not even suspect.”536 Elton made a more explicit exception for The Estate of Man by identifying its author and some actual content. That clue turned out to be worth following. More than a mere suggestion, Michael Roberts devoted a chapter to arguing that average intelligence in Britain (and elsewhere) was declining, and revisiting the familiar eugenic contention that fertility and intelligence are inversely proportional. He also described five intelligence levels, labeling them grades A through E, a ranking inevitably reminiscent of the Alpha through Epsilon castes of Aldous Huxley’s Brave New World.537,538 Roberts was unenthusiastic about the lower castes, musing, “It would be bad enough if any large section of humanity came to be regarded a sub-human inferiors, to be entertained and treated kindly, as an Englishman treats his dog, but not to be entrusted with any major decision. It would be worse, in the long run, for everybody, if millions of dull-witted machine-minders, film-fans and dog-race addicts were to tyrannize over their more intelligent fellow citizens … the right of the redundant, low-grade worker to 536 de Castro The Geography of Hunger (cit. n. 533), 295. 537 Roberts, The Estate of Man (cit. n. 534), 77-99. 538 Aldous Huxley, Brave New World (New York: Harper & Brothers, 1932). 240 go on living and reproducing his kind in a world to which he contributes far less than he receives … looms beyond the coming clash between rival unions each asking for a bigger share of the national income.”539 Although his concerns were wide-ranging, Roberts’ book also most directly foreshadowed Elton’s Invasions theme. In slightly under two pages (subheaded The Grand Ecology) he listed now-familiar environmental concerns that did not impinge on popular discourse for decades, including intercontinental transport of pest insects and disease organisms, biosecurity, unforeseen consequences of agricultural chemicals, junk food, traffic death rates, declining male fertility, interrupted fish migrations, reservoir silting, and drug-resistant disease. “It is not merely that each problem solved creates new problems,” Roberts wrote, “sometimes the solutions themselves are provisional and precarious.” He concluded: “We have made it possible for the world to become a single ecological unit, and we ourselves are desperately striving, through regulation, inspection, and regimentation, to save ourselves from the forces we have released.”540 As the Oxford-educated son of an Oxford-educated critic, and grandson of a “sometime Cambridge Fellow,” Charles Elton would identify with Roberts’ “A” caste, and it would be romantically anachronistic to suggest that he do otherwise. Elton made no effort to disclaim any aspect of Roberts’ views, including the eugenic defense of his caste. From such a perspective, any explosion of 539 Roberts, The Estate of Man (cit. n. 534), 132-133 540 Ibid., 102-104. 241 unwanted organisms, whether non-“A” humans or Colorado potato beetles, challenged the favorable status quo, evoked fears of lost control, and incited attempts (or at least thoughts of attempts) to reassert that control. In contrast with those (like de Castro) whose hopes for the hungry masses rested on the cheap, long distance redistribution of resources, Elton was convinced that the accumulating side effects of such distribution amounted not only to “ecological pandemonium,” but global ecological war. Unlike the higher profile World Wars, this one threatened shortages not only of commodities, but also of familiarity and predictability, rendering the world inhospitable and (he feared) possibly uninhabitable. But Elton’s view of the future was not uniformly gloomy. He felt that a livable world would inevitably require an instrumental relationship between “a modified kind of man and a modified kind of nature.”541 His hope for such conservation was buoyed by what seemed to be recent technological triumphs. Revisiting his opening “nuclear” analogy, and disclosing its purpose, Elton wrote: “Just now it is only possible to give a progress report and a hopeful forecast—the sort of thing a nuclear engineer might have given about power stations ten years ago.”542 Having begun a thread with atomic bombs, he ended it with (then) newly contained and “damped-down” nuclear power generation. Even after inflicting a casebook of horrors, Elton made clear that he was not wholly averse to intentional species introductions. While acknowledging that 541 Elton, Invasions (cit. n. 162), 145. 542 Ibid. 242 a direct ecological correlation of stability with complexity remained hypothetical, he recommended “the building up of fairly complex plant and animal communities:”543 “I believe that conservation should mean the keeping or putting in the landscape of the greatest possible ecological variety — in the world, in every continent or island, and so far as practicable in every district. And provided the native species have their place, I see no reason why the reconstitution of communities to make them rich and interesting and stable should not include a careful selection of exotic forms, especially as many of these are in any case going to arrive in due course and occupy some niche.”544 Though he wrote as an ecologist and lived a comfortably middle-class life, Elton’s avowed Invasion concerns were often proletarian. Given his concept of the ecological niche as a “profession,” Elton believed that the English natives were in danger of losing their jobs. He tolerated immigrants as long as they did not displace established locals. This attitude has familiar echoes in debates over human immigration, but its implications are ambiguous. In taking such a position, Elton would be endorsing the refrain that immigrants will occupy niches (that is, take jobs) that natives had refused or abandoned, and thus are economically—or in this case, ecologically— necessary. But he did not follow the corollary that empty niches might just as easily be considered opportunities for species with special “skills” such as high tolerance to salts or physical disturbance, and thus, like any skilled labor, worthy of a special investment. 543 Ibid., 151. 544 Ibid., 155. 243 The final pages touched on many aspects of (mostly British) conservation concern: retreating wilderness; inevitable human effects; the value of roadside hedgerows; tourism; timber; insecticide problems; and soil conservation. Elton invoked the “tripod” twice more, as if continuing to work it out. Previously he had appealed to religious motivation in terms of providing water for rats, then in creating refuges for wild species.545 In his last chapter he restated it as making “the countries we live in…safe for wild life.”546 The “aesthetic and intellectual” question became “making our surroundings interesting and satisfying,” then “more interesting.”547 The third, practical question metamorphosed from “land, crops, forests, water, sea fisheries, disease, and the like,” to “in order to promote stability of populations and a varied community in which all kinds of compensatory pressures will be exercised on populations,” to “more secure for the farmer, forester and fisherman.”548 His ultimate expression of the tripod was by far the most succinct: “refuge, beauty and interest, and security.”549 This was what Elton wanted from the world. The rest of the book described how the world was failing him, and whom he blamed.550 545 Ibid., 143; 151. 546 Ibid., 155. 547 Ibid., 144; 151; 155. 548 Ibid. 549 Ibid., 159. 550 Elton revisited and revised these ideas once more, in the closing pages of The Pattern of Animal Communities (cit. n. 382), 383; “The case for general conservation is threefold and has been discussed by me in an earlier book. It is moral … no power without responsibility. It is also based on interest, whether 244 Verging on completion, Elton seemingly paused to collect his thoughts: “The examples just given lead on to the idea of actually planning a better and more varied landscape.”551 He pleaded for traditional values. Once again, he turned to the common folk, for Hobbit-like wisdom: “In Britain we might still have the chance of keeping our own remarkable landscape before it loses its ecological variety. This landscape pattern was built up by individuals, mostly country people working by instinct and making a place to live in, not just a place to raise cash crops in.” Oxymoronically by American standards, he even proposed that this outlook “may enable us to put into the altered landscape some of the ecological features of wilderness.” 552 Elton prepared his readers for a closing rhetorical question by quoting John Muir (from Our National Parks, 1901) although the passage rather dubiously applied to a middling island nation: “‘To the sane and free it will hardly seem necessary to cross the continent in search of wild beauty, however easy the way, for they will find it in abundance wherever they chance to be.’ Will we be able to talk like this in fifty years’ time, as he could do fifty years ago?”553 personal, educational or purely scientific — the deep study of the world as it was when man found it, or those parts still keeping some of the character and potentialities of that earlier untamed richness … thirdly it is an economic question, a question of human survival, or at the very least man’s survival in environments that are worth surviving in.” In this version, explicit “aesthetics” have disappeared from the second “leg,” but begun tacitly to emerge in the third as “environments worth surviving in.” 551 Elton, Invasions (cit. n. 162), 158. 552 Ibid., 159. 553 Ibid. 245 War: the Dominant Motif Elton employed warfare explicitly as metaphor in his 1942 “big” book for ecologists (Voles Mice and Lemmings). There he reminded himself and his readers that “[anecdotal] accounts of these plagues abound in military metaphors, which testify to the serious threats that mice can offer to country people.”554 But Invasions was about nature being (mis)directed by human actions into a literal war against itself.555 Voles was written for scientists, and Invasions for everyone; reifying the metaphor, Elton finally cast his lot with the “country people” in 1958. Otherwise the title he proposed to Methuen, “The Ecology of Invasions,” seems rather weak, and his urgent, authoritarian calls to conduct further research and reform resource management would be idiosyncratic at best.556 It also seems likely that his publisher took the metaphorical view and added “by animals and plants” to clarify the matter. In Invasions, Elton identified himself as a partisan observer and commentator, rather than a participant. “I have described some of the successful invaders establishing themselves in a new land or sea, as a war correspondent might write a series of dispatches recounting the quiet infiltration of commando forces, the surprise attacks, the successive waves of later reinforcements after the first spearhead fails to get a foot hold, attack and counter attack, and the 554 Elton, Voles (cit. n. 247), 13. 555 Echoing Alfred Russell Wallace’s assertion of “actual war” in Island Life (cit. n. 135) (See Chapter One). 556 CSE to Samuel Abrams, Wheaton College, 7 Feb 1964. CSEP A28. 246 eventual expansion and occupation of territory from which they are unlikely to be ousted again.”557 Elton regularly substituted “enemy” for ecologically more precise terms, often giving prey “enemies” instead of predators.558 Variations of “invade” abounded. Even in the section emphasizing conservation philosophy, “invade” occurred thirteen times; it was accompanied there by seven “enemies,” two (each) “explodes” “penetrates” and “retreats,” and one (each) “attack,” “battle,” “conflict,” “conquer,” “threaten” and “thrust.” Most of those terms also recurred in other chapters, where species or populations also constituted “advance guards” and “bombardments,” established “bridgeheads,” encountered “chemical warfare,” “consolidated,” “dominated,” got “a firm hold,” kept “their forces intact,” were “mobilized,” “occupied” France, Belgium and Holland, “pressed” into new areas, “redeployed,” “seized [a] position,” “spread out from … headquarters,” accomplished “untold slaughter,” and “wiped out” other species. Analogies and Metaphors From the first to the penultimate page, bellicosity pervaded Invasions. Although Elton supplemented his military motif with “civilian” metaphors, many remained variations on the theme of competition. They played mostly supporting roles. Some of the major ones are discussed here. Elton almost imperceptibly framed his entire argument as an Atomic Age parable of “swords into ploughshares.”559 After his strong opening gambit 557 Elton, Invasions (cit. n. 162), 109. 558 Ibid., (e.g.,) 116; 146-7; 148; 150; 157. 559 Cf. Isaiah 2:4. 247 comparing ecological and nuclear explosions, he ignored the comparison for seven chapters, so that framing is faint and easy to overlook; it effectively became subsidiary to the general warfare motif. Perhaps the most interesting aspect of this idea is that it would become, at least in retrospect, deeply ironic. In “book three” Elton tacitly referred to Britain’s highly-touted Calder-Hall station at Sellafield, which in 1956 became the world’s first civilian source of nucleargenerated electrical power. It was then closely guarded knowledge that the Calder-Hall reactors were designed and built to produce weapons-grade plutonium. Furthermore, the first (1951) Sellafield reactors, called the Windscale Piles, were not only devoted to weaponry, but in October 1957 (between Elton’s submission of the Invasions manuscript and its publication) one of them experienced a core fire followed by a release of radioactive steam and gases that is still called “the West’s worst nuclear accident,” affecting not only Britain but continental Europe. It would be a quarter century before the incident was publicly acknowledged.560 Subsidiary versions of the control and containment analogy appeared throughout Elton’s discussion, most obviously wherever he appealed to the action or existence of “ecological barriers” (which he usually truncated to “barriers”) impeding the free movement of fauna and flora, or, as he put it, kept 560 Erik Martiniussen, Sellafield (Oslo: The Bellona Foundation, 2003) 30-31; Roger Highfield, “Cumbria still affected by fire at Windscale nuclear reactor 49 years on” http://www.telegraph.co.uk/ 25 April 2006. 248 them “cooped up.”561 Elton also mentioned confinement and obstruction, describing “Wallace’s Realms” as “colossal, separate nature reserves,” suggesting that containment worked both inwards and outwards.562 Elton invoked sports, recreation and games including chess, boxing, boating and hunting. His chess imagery suggests that he was not a particularly strategic player. “Man has [been] moving species around like chessmen” was not used in a context of thoughtful planning.563 Elsewhere, with similar scorn: “a good deal of chess play has also been done with clams.”564 Finally, Elton perhaps uniquely, contrasted playing chess with seamanship. “The world’s future has to be managed, but this management would not be like a game of chess— more like steering a boat.”565 This conception—what sort of boat, and under what conditions—is obscure. Elton seemed to be espousing a sort of command ecology. His call for a human hand on nature’s tiller suggested not only that nature needed guidance, but also that its direction could be improved. But although he had dutifully sailed as a passenger from England to Spitsbergen, Canada and the United States, Elton was not a celebrated boater. He was actually prone to seasickness.566 561 Elton, Invasions (cit. n. 162),, 34. 562 Ibid., 20, 31, 41. 563 Ibid., 33. 564 Ibid., 103. 565 Ibid., 151. 566 Southwood and Clarke “Charles Sutherland Elton.” (cit. n. 211) 249 According to Crowcroft, Elton even chose the name of his Bureau “while lying seasick in his bunk on a Cunard liner.”567 However puzzling it remains, the chess vs. boating analogy attracted at least one disciple. In 1982, well before the renaissance of both scientific and popular interest in Invasions, “social ecologist” Murray Bookchin quoted the passage in The Ecology of Freedom, calling it “a sensitive observation … what ecology, both natural and social, can hope to teach us is how to find the current and understand the stream. 568 But the analogy did not really catch on. Elton also wrote that humans had “knocked about” the world’s landscapes.569 In a slightly more developed analogy, “two species jostle” then “there is a very clear verdict … that hard fighting” has led to a particular outcome.570 Elton trained as a boxer during his undergraduate years. Providing an unlikely segue from boating, during the 1924 Oxford research expedition to Spitsbergen he engaged in a bout with Ian Colquhoun (later Rector of Glasgow University) on the deck of the 300 ton steamer Polar Bjorn after a rough crossing of the Barents Sea; Colquhoun challenged him with the declaration “no one ought to mind a bit of seasickness.”571 567 Crowcroft, Elton’s Ecologists (cit. n. 184), 12. 568 Excerpted in The Murray Bookchin Reader, Janet Biehl, Ed. (London: Cassell 1997), 35. 569 Elton, Invasions (cit. n. 162), 110. 570 Ibid., 122-123. 571 Charles Elton, unpublished manuscript, CSEP A33:24. George Binney, leader of this expedition, wrote in his book With Seaplane and Sledge in the Arctic (New York: George H. Doran Co.) that the Polar Bjorn “rolled from pure joy 250 Health related metaphors occur only rarely in Invasions. Perhaps the most straightforward concerns the introduction of a “counterpest,” or biological control organism: “Australia administered the poison, but it also supplied the antidote.”572 The remaining health metaphors are relatively minor. Elton referred to a few species besides Yersinia pestis as plagues (which might just as easily be categorized as a biblical allusion); and one aquatic plant “choked” a stream. Elton’s Data Universe I chose not to specifically identify most of Elton’s cases histories in the interest of describing his arguments rather than promoting them. He felt that these cases had some ecological feature in common: “[Invasions] are so frequent nowadays in every continent and island, and even in the oceans, that we need to understand what is causing them and try to arrive at some general viewpoint about the whole business.”573 He also maintained his conviction that they were deeply problematic: “The real thing is that we are living in a period of the world’s history when the mingling of thousands of kinds of organisms from different parts of the world is setting up terrific dislocations in nature.”574 Invasions mentioned by name (common or otherwise) at least 195 species that Elton considered foreign, human-facilitated, invaders.575 In summary: … alas for our landsmen, who lay prostrate in their bunks, their faces a pale green like ripening cheese.” 75 Elton, Invasions (cit. n. 162), 57-58. 573 Ibid., 18. 574 Ibid. 575 I may have missed one or two mentioned only vaguely or casually. 251 (indentation indicates a subcategory, but some of these are partial and do not “sum up”): Microorganisms: Pathogens: Diatoms and algae: 13 11 2 Vascular plants: Trees and shrubs: Grasses: Animals: Invertebrates: Worms: Mollusks: Arthropods: Crustaceans: Insects: Pests (USA): Vertebrates: Fishes: Salmonids: Amphibians: Reptiles: Birds: Mammals: Marsupials: Rodents: Ungulates: Carnivores: 15 7 2 167 114 4 15 95 15 72 47 53 15 7 0 0 16 22 1 7 9 2 This lists’s 10:1 ratio of animals to plants trivially confirms that Elton, although an avid gardener, was an animal ecologist, more familiar with the zoological than botanical literature. He and his Oxford colleagues worked mainly with rodents; but two-thirds of the animals he discussed in the book were invertebrates. Nearly two-thirds of these invertebrates were insects, and two thirds of the insects were pest cases from the United States (including Hawaii). Elton considered only one listed insect — the European honeybee — to be a 252 completely benign addition to the American fauna.576 As suggested previously, the predominance of American cases was no accident. The best-documented and most widely disseminated reports of animal invasion then available concerned problematic insects (domestic, agricultural, etc.) and their control. Americans were the vanguard regarding this issue. In the cases of two pest insects, Elton credulously repeated unlikely anecdotes of collection efficiency, if not actual abundance; and provided citations for neither. “By 1919,” he related, “a single person could gather up 20,000 [Japanese] beetles in a day.”577 Similarly, “When a bounty was offered for resting queens [of the wasp Vespula germanica] one schoolboy brought in 2,400 in less than a week!”578 Given in each instance a generous and uninterrupted ten-hour workday of actual collection, a beetle collector had slightly less than two seconds to acquire each insect; assuming “less than a week” to mean as many as six days, Elton’s intrepid schoolboy averaged ninety seconds to locate, recognize (as a queen) and capture each wasp. Elton’s willingness to propagate such hyperbolic claims did not bode well for his general credence, but there is no evidence that anyone ever objected to them. With a few regionally notorious exceptions, invading vertebrates were not yet a very public concern by the 1950s. Few members of Elton’s audience 576 “Africanized” honeybees first escaped research quarantine in Brazil in 1957; their survival and potential extent of their spread was not evident until much later. 577 Elton, Invasions (cit. n. 162), 52. 578 Ibid., 92. 253 needed reminding that rats were a problem, but rat populations were concentrated in urban sewers, around docks and on farms, where relatively few of his likeliest readers would encounter them. Escalating British and European populations of (once) American muskrats and grey squirrels had been among Elton’s favorite whipping boys since the 1920s, but he rarely pressed the issues publicly. The British, unlike their continental counterparts, had managed to eradicate muskrats well before World War Two, and anti-squirrel sentiment was patchy. Feral domestic pets like stray dogs and cats were categorically difficult to fold into the invasion mix (although, as we saw in Chapter Two, Aldo Leopold willingly took on cats.) For vertebrate cases, Elton relied mainly on introduced game animals (four species of deer and two upland game birds), and sport fishes (including seven kinds of salmon and trout). Proper hunting and fishing, as opposed to poaching, were traditionally reserved for the British gentry. The advent of these species, then, had more obvious advantages than disadvantages to socially privileged sportsmen, and probably didn’t matter, in a proximate way, to anyone else. The absence of amphibians and reptiles among Elton’s examples is surprising. By 1955 there were already growing and well-documented populations of at least two introduced frogs in Britain, and indications were that at 254 least one introduced lizard would persist in southern England.579 Later global “poster children” of herpetological neobiota were already on the move as well, but their case histories would not be compiled for another two or three decades. Central American cane toads (Bufo marinus) were island hopping at the behest of Caribbean sugar farmers before 1900 and across the South Pacific by the 1930s.580 Brown tree snakes (Boiga irregularis) known previously from northern Australia, New Guinea and surrounding islands were reported on Guam by the early 1950s, but did not become a cause célèbre until they began interrupting electrical service late in the 1970s.581 Plates and Figures as Facts Invasions included an eclectic assortment of fifty-one (including the frontispiece) black-and-white plates. One was a map. One was a line drawing. Thirty-nine were photos, ten by the author. Four of these ten were dated 1957, and may have been shot specifically to illustrate the book. Fourteen photos showed landscapes or vegetation without significant foreground features. Five photos focused on people doing something, and several more included humans in the frame, apparently for scale.582 Fifteen photos depicted specific organisms. 579 Christopher Lever, Naturalized Reptiles and Amphibians of the World (Oxford University Press, 2003) 106-107, 218-219, 222-223. 580 Ibid., 147-175. 581 Ibid., 125-133. 582 Elton’s wife Joy Scovell provided scale in Plate 40, as (most likely) did his son in Plate 45. 255 Twelve plates reproduced paintings. Five of these depicted specific organisms, one a cultural feature, and the remaining six were romantic, dioramalike assemblages of animals originally painted to illustrate Alfred Russell Wallace’s 1876 The Geographical Distribution of Animals. These, like the “balance” metaphor, will be discussed in the final chapter. There were also fifty-one text figures, all appearing in chapters one through seven (i.e., in the first two books of the “trilogy”). Forty-eight of them were, or included, maps. Two depicted organisms (a seaweed and an insect) and the other was Bruegel’s drawing. Unlike the plates, all of the figures were reproduced from other publications. Doubtless there were “losses in translation” from original context to Invasions, and those losses vary, from the fundamental to the negligible. The “feeling” Elton needed his figures to convey was a sense of change: the presence of a species or disease in a “new” location and (ideally) an enlarging affected area. As a result, many of them involve time-series, showing advancing frontiers or expanding polygons. This visual simplification helped Elton reduce the multifarious phenomena under discussion to variations on the militaristic theme of territorial conquest and occupation. As such, the figures should be—and are—reminiscent of wartime invasion front maps. Figure 4.3 compares a map from a 1920 book about World War One, a 1928 USDA map reproduced for Invasions, and a July 1944 map of German advances and retreats on the Eastern Front. 256 Figure 4.4.a. “The German Invasion of France” from a 1920 account of World War One. Note particularly the two successive Allied Lines, labeled 21 August and 5 September 1914.583 583 A. F. Pollard, A Short History of the Great War (London: Methuen & Co., 1920). 257 Figure 4.4.b. “Breeding range of the European starling in eastern North America” from a 1928 USDA circular. Elton re-captioned this to use Invasions Figure 5, with typeset years replacing the hand lettering. 584 584 May T. Cooke The Spread of the European Starling in North America (United States Department of Agriculture, Circular No. 40, 1928) accessed online 4September 2006 at http://www.wku.edu/~smithch/biogeog/COOK1928.htm. 258 Figure 4.4.c. “The German Catastrophe in White Russia” (1944) from The Illustrated London News shows another distinct time series of advancing (and in this case, retreating) fronts.585 Elton seems to have treated maps themselves as tantamount to data, rather than as documents representing data. By borrowing maps from numerous previous publications he saved time and production costs, but glossed over variations in their three basic, manipulable attributes: scale, projection, and 585 Cyril Falls, “The Great World War: Fronts — Western and Eastern.” Illustrated London News (8 July 1944) 46. Map “copyright by ‘The Times’ (undated). 259 symbolization.586 Only seven of the maps he used have identified scales. None identifies its projection. Symbolization varies widely. Numerous ways to represent “occupied” or affected territory appear. There is no consistency to relative sizes of dots on the six different point distribution maps, and a sort of pointillist shading on two more maps suggests similar significance without carrying it. Even reproduction quality varies significantly, leaving some figures, including Elton’s first two practically illegible. These, in particular are problematic for several reasons. Elton’s figures one and two were each half-page reproductions, approximately 70% of original scale, of unnumbered illustrations appearing among the appendices on pages 104 and 111 of the 1955 U.S. Public Health Service (PHS) monograph, A History of Plague in the United States. Both are maps of (then, all) forty-eight states, with state borders outlined, and information on the incidence of plague (Yersinia pestis) represented by shaded areas. The units of representation are counties. This is administratively relevant for public health purposes; but counties obviously do not contract the disease. Nor are they strictly comparable. For example, counties in California vary in land area by well over two orders of magnitude (from San Francisco County at 47 square miles to San Bernardino County at 20,053 square miles).587 California is a most 586 Mark Monmonier, How to Lie with Maps (Chicago: University of Chicago Press, 1996) 5. 587 U.S. Census 2000 Tiger file data, rounded to the nearest square mile. It was assumed that the areas of these counties did not significantly change between 1955 and 2000. 260 germane example, since all but a very few of the counties shaded in each map lie in the western half of the country (i.e., west of 100° west longitude). They are not reproduced here because even with originals in hand, the prospect of their adequate reproduction is low). In the PHS document, the map that became Elton’s Figure One was titled “Counties in the United States where human plague has occurred, 19001951.”588 It accompanied a table listing affected counties and the number of plague cases and attributed deaths recorded in each one.589 Thirty-six counties were shaded. Thirty of those were in the west. Nineteen were in California, including San Bernardino and San Francisco. According to the table, sprawling but (then) almost unpopulated San Bernardino County, physically larger than several entire eastern states, had only one recorded case of plague and no attributed deaths, the lowest possible figure that could be represented. Virtually invisible San Francisco County had 289 cases and 196 deaths; the highest recorded in the table. For his purposes, Elton re-titled the map “Counties in the United States where plague has occurred in man,” a reasonable approximation of (but unnecessary substitute for) the original.590 His referring text, “In the United States and Canada a similar underworld of plague (with different [animal] species 588 Vernon B. Link, A History of Plague in the United States of America, Public Health Monograph 26 (Washington DC: U.S. Department of Health, Education and Welfare 1955), 104. 589 Ibid. 590 Elton, Invasions (cit. n. 162), 16. 261 in it) is established over an immense extent of the Western regions” conveys no sense of the relationship between populations and areas in the pictured region— unexpected indeed for an author preoccupied with populations, areas and regions.591 Elton’s Figure Two had additional problems. Originally titled “Counties in the United States surveyed for animal plague, 1900-1950,” it was distinguishably—but not very clearly—shaded in two densities, one indicating negative survey results and the other positive results.592 When reproduced, the already problematic shading of the “negative” counties became hopelessly indistinct.593 Under Elton’s title, “Counties in the United States where plague has occurred in rodents,” the negative/positive distinction was unexplained, and became worse than meaningless.594 The new title itself was also misleading. The unnumbered table accompanying the original PHS map distinguished between two classes of “animals examined”: “domestic rodents” and “wild animals.”595 Table Six in the body of the PHS text made it clear that even though plague was generally associated with (unspecified types of) ground squirrels, animals other than 591 Ibid., 17. 592 Link, A History of Plague (cit. n. 588), 111. 593 Elton, Invasions (cit. n. 162), 17. Both the 1958 (original, Methuen) and 2000 (facsimile, University of Chicago Press) editions of the book were examined. The 1958 original was considerably more distinct, but not usefully so. 594 Ibid. 595 Link, A History of Plague (cit. n. 588), 110. 262 rodents (rabbits, weasels, badgers, moles, “wildcats,” house cats, and coyotes) were also tested during some surveys.596 Furthermore, a full-page listing in the PHS Appendix of “Animal sources of plague-positive ectoparasites, by state” is populated mostly by rodents, but also includes a rabbit, a hare, badgers, and even burrowing owls.597 That list, and the methodological description in the main PHS text indicated that both PHS and Elton should have emphasized that the data used in preparing this map came from several studies that, over time, involved different methods for determining the presence of plague bacillus. Some studies focused on identifying infected animals, and others on ectoparasite vectors of the bacillus. British (and even most American) readers unfamiliar with the vagaries of western U.S. political geography could hope find little real meaning in either set of blotches. Regardless of their applicability to public health administration, the demographic ecological and epidemiological precision of these maps was practically nil; and for most of the audience, consulting the original source was out of the question. The idea of marking one non-fatal, human case of plague in a desert region over twice the size of Wales more prominently than nearly 200 human plague deaths within a major city might understandably have surprised many in Elton’s audience. All the unwary non-specialist could hope to glean from these figures was a sense that the western U.S. (including Hollywood!) was rife with plague, and wonder why this hadn’t made headlines. 596 Ibid,. 30. 597 Ibid., 113. 263 It cannot be said for certain that Elton deliberately misrepresented the meanings of these maps. But the available explanations boil down to a choice of prevarication or carelessness. Once again, Elton’s emphasis on speedy composition may have had an impact. It is also possible that personal recollections of his 1938 visit to California, which included a tour of the plague field survey “near San Francisco” led him to assume that what he recalled of his experience there was representative of the entire effort.598 Either way, his conception, analysis and presentation of the plague phenomenon in North America suffered as a result. Early Reactions Some of Elton’s contemporaries seemed convinced by his proposition that variety, or species diversity, conferred ecological stability: “Without doubt, there is a case for accepting Elton’s stimulating hypothesis. Still more may we follow him in his plea for the conservation of diversity in habitats modified by man.”599 This idea still has proponents and opponents, but has never been shown have the kind of practical application Elton hoped for. The relatively few reviewers of Invasions found “book three,” regarding conservation philosophy, much more compelling than the extended litany of cases that composed the bulk of the work. One wrote: “in the last two chapters [Elton] deals with ‘The Reasons for Conservation’ and ‘The Conservation of 598 Elton, Invasions (cit. n. 162), plates 1,2 and 3 (unnumbered pages following 32). 599 W.H. Pearsall, “The Ecology of Invasion: Ecological Stability and Instability,” New Biology 29 (1959), 95-101. 264 Variety’. The earlier chapters are, in a sense, no more than an elaborate introduction to these two, which for one reader at least are the most original and significant part of the book.”600 Another expanded: “The rest of the book would have been brought into sharper focus if these last two chapters had preceded the others. They serve as a general statement of Elton’s thesis: that ‘conservation of variety’ is preferable to a simplification of the ecological community, on three counts…(1) there is a ‘right’ relation between man and animals, from a religious point of view; (2) there would be a better opportunity for a richer experience of observing nature; and (3) ecological stability would be promoted and disastrous ecological ‘outbreaks’ prevented.”601 A third summarized: “At the very end, the author suggests that, prior to changing, cutting away, draining or spraying an area, it is vital to ask three questions: (1) What animals and plants live in it? (2) What beauty and interest may be lost? (3) What extra risk will changing it add to the accumulating instability of communities?”602 It seems that these readers, at least, did not wholly appreciate the brute force repetition Elton had provided. It also appears that none of them were familiar enough with A Sand County 600 P.W. Richards, [Review of] Elton, C.S. “The Ecology of Invasions by Animals and Plants” The Journal of Ecology 47 (1959), 525-526. 601 Margaret Foreman Cohn, [Review] “The Ecology of Invasions by Animals and Plants” by Charles S. Elton.” The Quarterly Review of Biology 34 (1959), 303304. 602 Walter P. Taylor [Review] “The Ecology of Invasions.” (cit. n. 219). 265 Almanac to comment on the similarity between the Eltonian and Leopoldian views, but wide acknowledgement of the latter was still several years away.603 Elton’s views on conservation encouraged another proto-environmentalist writer. After a young Edward O. Wilson recommended Invasions to her, Rachel Carson, who was then (1958) compiling Silent Spring, replied: “I found [Invasions] enormously stimulating. It cuts through all the foggy discussions of insect pests and their control like a keen north wind.”604 Carson’s British publisher Hamish Hamilton later asked Elton to preface the British edition of Silent Spring, but he begged off, writing to Carson “I explained to Mr. Hamilton that I think it is best for authors to preface their own books, especially when the subject is such a controversial one as this,” and apologizing for his inadequate knowledge of toxic chemistry: “I would not feel able to answer the questions that would be shot at me.”605 Carson replied, “I completely understand your reasons … may I take this opportunity to tell you how much your various books have meant to me over the years? Your recent Ecology of Invasions was especially fine and I thought it did 603 Meine, Aldo Leopold (cit. n. 163), 526. 604 Rachel Carson to Edward O. Wilson 6 November 1958. CSEP A.72. Wilson forwarded a copy of this letter to Elton with an undated cover memo: “While searching through my correspondence files the other day I came upon this letter, which I thought might interest you personally. Under the circumstances [unspecified] I felt it would be all right to send you a copy.” 605 C.S. Elton to Rachel Carson, 10 September 1962. CSEP A.72. Wilson’s recommendation to Carson was mentioned in Linda Lear’s biography Rachel Carson: Witness for Nature (New York: Henry Holt, 1997) 332-333; Carson’s request to Elton and his refusal were omitted, although the practically concurrent discussion with Lord Shackleton was mentioned (Ibid., 422). 266 much to set in proper perspective the problem I have dealt with in Silent Spring.”606 The first British edition of Carson’s classic ultimately appeared with an introduction by Lord [Edward] Shackleton and a preface by Julian Huxley. Elton’s reticence may thus have cost Invasions some additional attention, but he apparently considered edging into the limelight on Carson’s behalf too high a price to pay for reinforcing their obviously mutual concerns. An early formal citing of Invasions, and perhaps even the first, came in G. Evelyn Hutchinson’s 30 December 1958 Presidential Address to the American Society of Naturalists, famously titled “Homage to Santa Rosalia, or Why are there So Many Kinds of Animals?” Hutchinson was polite, if not enthusiastic, and perhaps a bit mischievous. He called Invasions “a fascinating work largely devoted to the fate of species accidentally or purposefully introduced by man.” 607 This played havoc with Elton’s actual sympathies, while broaching (without answering) a potential ethical question Elton had not addressed: what duties are owed to introduced species? Hutchinson, who had then recently proposed a niche concept to supersede Elton’s, also took aim at the possible shortcomings of other major metaphors associated with Elton’s work (e.g., food chains and the pyramid of numbers.) Elton probably would not have enjoyed the presentation. 606 Rachel Carson to C.S. Elton, 24 September 1962. CSEP A.72. Incidentally, Lear (Rachel Carson, 521, n.6) concluded that Carson had never read Aldo Leopold’s A Sand County Almanac. 607 G. Evelyn Hutchinson, “Homage to Santa Rosalia, or Why are there So Many Kinds of Animals?” The American Naturalist 93 (1959) 145-159. Perhaps, since it was revised and published later, the mention was added after the fact. 267 Elton’s 1958 proof copy of Invasions survives among his papers in the Oxford Bodleian Library. Perhaps considering a revised edition, he had crammed it with news clippings and notes dating between 1958 and 1986, “filing” them between the pages mentioning the taxa concerned. A list of addenda included (Marston Bates’ adviser) Frank “Egler on herbicides & management of US ‘utility strips’ and roadsides.” Also preserved among the associated materials is Elton’s abstract of Edward O. Wilson’s March 1958 Scientific American article “The Fire Ant.”608 After the Fact In the years following its Methuen/Wiley debut, Invasions was republished several times in facsimile editions: in London by Methuen (1963, 1966), by the English Language Book Society (1966, 1972) and by Chapman and Hall (1977, 1982); in the United States by the Halstead Press (1977) and University of Chicago Press, 2001). It was translated into Polish by Zdzisława Wûjcik and published in Warsaw (1967) as Ekologia inwazji zwierząt i roślin. In his unpublished memoirs, Elton noted with satisfaction “This book was well reviewed and was reprinted several times. It is still on sale (1989) now as a paper-back edition.”609 608 CSEP A.28; Edward O. Wilson, “The Fire Ant” Scientific American 198 (1958) 3:36-41 609 Charles Elton, unpublished ms. CSEP A.33:43 268 He also proclaimed, “Little notice had been taken of this very important subject until my book came out.”610 This assessment is difficult to credit, since Elton managed to cite nearly 200 published case studies and included nearly 300 references in Invasions. Even as author 301, Elton apparently remained ignorant of much potentially relevant literature, most particularly from phytogeography. This hazard is familiar to anyone compiling a supposedly comprehensive review. It is particularly ironic in light of a rather severe comment Elton made shortly before he embarked on the Invasions project, when evaluating an Oxford colleague: “As you don’t read much, you will overlook your predecessors.”611 He did not warn about ignoring or dismissing them unaddressed. It is also hard to credit his apparent decision to omit the opinions of authors demonstrating moderate views (like Bates), and significant “faunal histories” (like Ritchie’s The Influence of Man on Animal Life in Scotland) he had cited in his own previous works. It cannot be said that Elton didn’t read much. He used up many pages in The Journal of Animal Ecology with annotated lists of recent papers and books. Elton also spent a great deal of time collecting and cataloging data. But despite a lifetime of fieldwork, his empirical knowledge of “invasions” extended to only a handful of cases, and he was not the primary authority regarding any of them. 610 611 Ibid. C.S. Elton to Dennis Chitty (ca. 1956) quoted in Chitty, Do Lemmings Commit Suicide? (cit. n. 187), 122. 269 Indeed, since he rarely left Britain, his knowledge of most of the ecosystems discussed in Invasions was entirely derivative. To stand as a seminal contribution, Elton’s review needed to be something special indeed. He clearly believed he had made the topic his own. But Elton was never comfortable as an activist, and by the time Invasions was published he had withdrawn from any formal role in shaping British conservation policy. In contrast with Rachel Carson, he settled for describing battles rather than fighting them. And in contrast with both Carson and Aldo Leopold, he survived in (public) obscurity long after his book’s publication. But despite his best efforts to remain “private,” he could not escape posthumous celebrity. Like Leopold and Carson, he was remade as a culture hero. Ironically, his combination of personal obscurity and professional celebrity were perfect fodder for mythmaking. Eventually a new cadre appropriated him as their doctrinal founder, and hoisted his standard over their boot camp of militant eco-nativism. Chapter 5: Elton and Invasions, Reconstituted and Retired The Ecology of Invasions by Animals and Plants rested in “cult classic” semi-obscurity for decades. Charles Elton did not set aside his Wytham Wood research in order to study (or defend against) “invasions.” He made no apparent effort to organize his colleagues into a new ecological subdiscipline, or to enlist Oxford ecology students to study the phenomena he described in his book. It was as if Elton had emerged from a cloister to preach a single sermon, then retreated to a life of contemplation. The Ecology of Invasions was cited occasionally, but not inevitably, in the books and papers about human redistribution of biota that began appearing in the 1980s. One of the earliest major efforts, a conference proceedings titled The Ecology and Management of Biological Invasions in Southern Africa, included twenty-five papers by a total of fifty authors, but only one of those papers cited Elton’s Invasions.612 A few years later, half of the chapters in Biological Invasions: A Global Perspective cited Elton’s book.613 Although it is difficult to determine the exact number of publications about human redistribution of biota in the 1990s, a citation search suggested that it exceeded 300; but only nine of them cited The Ecology of Invasions by Animals and Plants. Elton directly informed only a fraction of invasion biology’s overall 612 I.A.W, Macdonald, et al, Ecology and Management of Biological Invasions in Southern Africa. (cit. n. 2). 613 J. A. Drake, et al, eds. Biological Invasions: A global perspective. Vol. 37, SCOPE. (Chichester: John Wiley and Sons, 1989). 271 output during the 1980s and 1990s. Both the overall output and the number of citations exploded after 2000; from 2000 to the latest indexed publications (partial 2006), The Ecology of Invasions was cited seventy times.614 Invasion biologists had a different impression. In 2001, Davis, et al wrote, “The unofficial father of invasion ecology is Charles Elton, whose seminal 1958 book … has been cited by virtually every major invasion paper since its publication.”615 Petr Pyšek was less circumspect: “in the current world biological invasions have ceased to be the domain of a small band of scientists who recognized the magnitude of the problem long before their colleagues and the public. The father of invasion ecology, Charles Elton, who made historical steps in his 1933 London Times article and ‘Balance and Barrier’ broadcast in 1957 is probably the most outstanding example of such a visionary.”616 On reflection, Davis later revised his estimate. “Elton was not the father of invasion ecology, but he might be considered the founder of one path of invasion ecology. This path, the conservation and environmental path, has been well traveled and maintained during past several decades.”617 614 ISI Web of Science queries, 9 November 2006. 615 Mark A. Davis, Ken Thompson, and J. Philip Grime “Charles S. Elton and the Dissociation of Invasion Ecology from the Rest of Ecology” Diversity and Distributions 7 (2001) 97-102. 616 Petr Pyšek “On the Road to Understanding: From John Wyndham To Reality” Diversity and Distributions 9 (2003) 251–252. 617 Mark A. Davis. “Invasion Biology 1958-2004: The Pursuit of Science and Conservation” in M. W. Cadotte, S. M. McMahon, T. Fukami, eds.. Conceptual Ecology and Invasions Biology: Reciprocal Approaches to Nature. (Dordrecht: Springer 2005) 35-64. 272 Invasion biologists occasionally acknowledge pre-Eltonian authors, but dismissively. A recent example: “In the 19th century, Alphonse de Candolle, Charles Darwin, and Joseph Hooker already paid a substantial attention to plant invasions. However, only after the publication of Charles Elton’s (1958) classic The Ecology of Invasions by Animals and Plants did invasion ecology emerge as a new discipline, first slowly … and later explosively.”618 It was if anything that happened before 1958 simply did not or could not really matter anymore. Concurrently with this project have come tentative signs of change. At the 2006 annual meeting of the Ecological Society of America, James Carlton gave a talk identifying a number of pre-Invasions books, beginning (chronologically) with German cultural historian Victor Hehn’s 1885 Wanderings of Plants and Animals from their First Home. Also in 2006, a major academic publisher inaugurated a “series in invasion ecology” with an edited volume including a chapter titled “Darwin to Elton: Early Ecology and the Problem of Invasive Species.”619 Following that chapter came “Invasion Biology 1958-2005: The Pursuit of a Science of Conservation.”620 As their titles indicated, the former pushed the discussion back to Darwin, specifically, to chapter three of the first edition of The Origin of Species (1859), and closed with Elton’s “magisterial book on invasion 618 Marcel Rejmánek, “Invasive Plants: What We Know and What We Want to Know” American Journal of Botany 92 (2005) 901–2. 619 Marc William Cadotte, “Darwin to Elton: Early Ecology and the Problems of Invasive Species,” IN Cadotte, et al, eds. Conceptual Ecology and Invasion Biology (cit. n. 617), 15-33. 620 Mark A. Davis, “Invasion Biology 1958-2005: The Pursuit of a Science of Conservation.” IN Cadotte, et al, eds. Conceptual Ecology and Invasion Biology (cit. n. 617) 35-64. 273 biology.”621 The latter began with “Charles Elton’s invasion classic” and ended, chronologically, with the 2004 meeting of the American Institute of Biological Sciences.622 Strangely, aside from those statements, Elton’s Invasions was a lacuna. Neither author attempted to explain the book or its significance. More puzzling still, the series editor’s foreword laconically and oddly called Invasions “Elton’s delightful if not foreboding treatise,” following with “of course Elton’s assessment was correct and indeed the problem is even greater today.”623 These statements suggest that, whatever else can be said about the history of this topic, the advent of Invasions plays a pivotal, perhaps ineffable role in invasion biology. In 2000, after William Faulkner scholar and occasional (best-selling) science writer David Quammen called it “one of the central scientific books of our century,” the University of Chicago Press issued a facsimile reprint of The Ecology of Invasions by Animals and Plants.624 This stimulated a new round of review and comment, providing an opportunity to examine the book in a new context. The edition included a new 2,500-word “Foreword” by Daniel Simberloff, a Harvard protégé of Edward O. Wilson, now (2006) Nancy Gore Hunger Professor of Environmental Science at the University of Tennessee. First among his specific research interests, Simberloff lists “Invasion biology[:] The patterns 621 Cadotte, “Darwin to Elton” (cit. n. 619), 15-17. 622 Davis, “Invasion Biology 1958-2005” (cit. n. 620), 50. 623 James Drake, “Foreword” to Cadotte, et al, eds., Conceptual Ecology and Invasion Biology (see cit. n. 617), xv-xvi. 624 Christie Henry, Senior Editor, University of Chicago Press, to M. Chew, 10 February 2004. 274 displayed by species introduced outside their geographic ranges, the impacts such species have on the communities they invade, and the means by which such invasions can be managed.”625 Simberloff’s “Foreword” to Invasions, more than any other extant work, exemplifies the way invasion biologists claimed Elton as their disciplinary father, and how Elton’s work was recast in the process.626 The anachronistic grafting of a late 1990s environmentalist’s heading onto a body of mostly 1930s-era conservation ideas had a pretty definite purpose. Simberloff’s “Foreword” made Elton the father of invasion biology. It taught a new audience how to read the significantly outdated Ecology of Invasions. It described which of Elton’s assertions to celebrate, and which to forgive and forget. And in the process of this re-visioning, Simberloff carved a niche for himself as Elton’s vicar-successor, and as a defender of anti-invasion dogma. Simberloff began: “Charles Elton was a founder of ecology.”627 Historians of ecology (and Elton himself) disagreed with this assessment of Elton’s role.628 After mentioning his 1927 Animal Ecology, Simberloff called Elton’s 1933 letter to the Times “his first publication on [invasions by introduced species];” but Animal Ecology itself demonstrated otherwise. What Simberloff saw in Elton as an “early 625 Daniel Simberloff, http://eeb.bio.utk.edu/simberloff.asp accessed 21 October 2006. 626 Although this makes Simberloff the unfortunate (though hardly unwitting) target of my analysis, his introduction of Invasions is an archetypal apologia for invasion biology. 627 Daniel Simberloff, “Foreword” in Charles Elton, The Ecology of Invasions by Animals and Plants (Chicago: University of Chicago Press 2000). 628 Robert P. McIntosh, The Background of Ecology. (cit. n. 218) 61-62, Donald Worster, Nature’s Economy (Cambridge: University Press,1994), 294-5. 275 concern with invasions by introduced species” [emphasis mine] trailed half a century behind Wallace’s, and further still behind Hooker’s regarding St. Helena. Simberloff called The Ecology of Invasions “the book that founded a whole field of research,” but no matter how narrowly construed, that claim seems unsupportable. Historical accuracy was obviously not a motivation, which is somewhat baffling in the context of invasion biology’s obsession with habitations, that is, with what even Elton called “accidents of history.” But with those opening bluffs, Simberloff simply declared all the historical justification that invasion biology required, and assumed the responsibility of staking its claim to a classic, but still modern-ish, scientific pedigree. “Yet,” he continued, “the book is aimed at a lay audience.” Simberloff left hanging the reasons that a book intended to found a specialized field of research would be formulated for tyros. He failed to consider that Elton might have written Invasions mainly for self-satisfaction, and by association to satisfy a waning population of “scientific naturalists,” the last vestiges of an all-but-extinguished nineteenth-century tradition. On that basis, Invasions marked a disciplinary conclusion, not an inception. Elton saw Alfred Russell Wallace as a colleague in a way Simberloff could never see Elton. The quiescent years between 1958 and 1982 were more than a hiatus, or a period of incubation. They marked another episode of the theoretical exhaustions and empirical recapitulations that have characterized the study of human redistributions of biota for over two centuries. Once all that could be said was said, the study was laid aside to again await rediscovery. 276 “The science of invasion biology has three basic components” wrote Simberloff, “and, though previous authors had recognized some of them, Elton was the first to unite them and recognize their interdependence … first, over hundreds of millions of years, the plant and animal communities of the different continents have come to be very distinct from one another. Second, human trade and travel are rapidly obliterating these distinctions. Third, the process has grave implications for the conservation of diversity.”629 Elton explicitly claimed to be drawing on “faunal history,” “ecology,” and “conservation.”630 Simberloff’s first component mirrored Elton’s own, as did his third, albeit anachronistically. Elton was concerned with conserving variety — “keeping or putting in the landscape the greatest possible ecological variety” — because he believed (but could never show) that ecological variety was the key to the practical goal of conferring ecological stability. But that motivation is a far cry from conserving the metaphysically expansive “biodiversity” sensu 1999. Later in the piece Simberloff dismissed Elton’s belief that “diversity” conferred a practical and desirable ecological stability, again substituting his own terminology and concepts for Elton’s, without explanation or acknowledgement.631 But he had explained this before: “Some of my colleagues will argue that biodiversity is an instrument to human welfare, even if they don’t believe it. … I won’t. I won’t make that argument. My real temptation is to say that if a person doesn’t know the value of a species, even if there’s no practical 629 Simberloff, “Introduction” (cit. n. 627) vii-viii. 630 Elton, Invasions (cit. n. 162), 13. 631 Simberloff, “Foreword” (cit. n. 627) xii. 277 or economic concern, then I can’t say anything that will convince that person. It would really be something like a religious conversion. It’s essentially a moral or ethical kind of thing.”632 For Simberloff, it seems, Elton’s “third question,” the one “hanging over the whole world,” was not a source of gravitas. Yet in the end he relented, because invasion biology’s own relevance and future (including his own) hung in the balance: “there is still much for invasion biologists to do…just as [Elton] predicted, the practical consequences of this problem have grown enormously.”633 Elton warned, but was also resigned to the fact, of ongoing, unintended biogeographic outcomes as a result of rapid, long-distance transportation. Simberloff’s anomalous “second component” substitution played up this theme, the “breaking down of Wallace’s Realms,” but simplified it in the extreme. The blanket indictment, “human trade and travel are rapidly obliterating [evolved] distinctions [between continental biotas]” sans any acknowledgement of the benefits of trade and travel (e.g., educating, provisioning and relocating invasion biologists) suggests, at minimum, the elite obliviousness of the ecotourist. The ultimate purpose of trade and travel is the breaking down of barriers. It seems incoherent to simultaneously decry and exploit that effect, or to exploit it in order to decry it. Global, bulk and mass transportation amount to a whole new set of gradients along which any fortuitously positioned and adapted individual or 632 Carol Kaesuk Yoon, “Do Earth’s Species Face Death?” Oregonian (15 August 1991) E1-2; David Takacs, The Idea of Biodiversity: Philosophies of Paradise (Baltimore, Johns Hopkins University Press) 335. 633 Simberloff, “Foreword” (cit. n. 627) xiii. 278 propagule (egg, seed, spore, etc.) might be dispersed. The biosphere is no longer a sphere at all, because its new topology connects previously unconnected points on its “surface” in a unprecedented (but non-random) way. The new biospace must be mapped at multiple concurrent scales to account for the speed and commonness of travel between locations, and even for the tonnage of freight in transit. It’s as if the old surface had been folded and refolded, bringing formerly distant locations face to face. For example, all major commercial ports have intermittently come in essentially direct contact at decreasing intervals for the past 150 years (some for centuries longer). They will do so into the foreseeable future. As Thellung’s inventory around Port Juvenal and Carlton’s in San Francisco Bay show, the history of commerce is now recorded in the biota of those habitations. Charles Elton pointed out that quarantines have limited capacity to resist such an overwhelming potential. Perhaps, then, invasion biology’s fundamental motivation is to perfect humanity by disembodying it, whatever that might entail; only non-corporeal beings without material needs can avoiding unintended physical outcomes, and thus meaningfully aspire to the ethic of “leaving no trace.” How better to prevent humans from influencing ecology than by rendering them non-ecological? Simberloff emulated Elton’s strategy of listing case after case of “horrors.”634 The identities of the species remain less interesting than the terms used to describe them. Invasion biology’s obsession with containment was in full 634 Ibid., ix. 279 view, and it took an interesting form. Unlike habitats, regions must be arbitrarily defined. Thus Simberloff claimed that species have invaded political entities like California, England, Louisiana and Chicago, or whole continents, or vague territories like the Pacific Northwest. His invaders are “German” wasps, “African” snails, “Chinese” crabs and “European” mussels. None of these are ecologically descriptive. Most are political constituencies, administrative powers, regional identities and citizenships: in short, they return us to Watson’s “civil claims.” The formula is a simple one, and boils down to a common complaint: your dog is in my yard! By extension, then, invasion biology aspires to the office of planetary dogcatcher. Charles Elton was a cautious optimist regarding the potential for using populations to damp down one another’s dynamics, including the introduction of “counterpests” or biological control organisms. Simberloff, on record since 1991 as a pessimist in this regard, wrote “Elton discussed some successes and failures of biological control,” but then proceeded to list only the failures.635 Simberloff also tried his hand at Eltonian rhetoric, calling the failure to suppress a target species accompanied by the accidental extinctions of “at least 30 species and subspecies of endemic snails on Pacific islands, a hecatomb rarely if ever 635 Simberloff, “Foreword” (cit. n. 627) ix; Daniel Simberloff, “Keystone species and community effects of biological introductions. IN Assessing Ecological Risks of Biotechnology, ed. L. R. Ginzburg. (Boston: Butterworth-Heinemann, 1991) pp. 1-19; Simberloff, D. (1992). “Conservation of pristine habitats and unintended effects of biological control.” In Selection Criteria and Ecological Consequences of Importing Natural Enemies, ed. W. C. Kauffman & J. E. Nechols. (Lanham, MD: Entomological Society of America)103-117; Daniel Simberloff and Peter Stiling, “Risks of Species Introduced for Biological Control,” Biological Conservation 78 (1996)185-192. 280 matched by a single deliberate human action.”636 This was a stumble; hecatomb sacrifices were deliberate killings, but the “non-target” extinctions he referred to were neither deliberate nor foreseen. And the extinctions were clearly regrettable even to those who unwittingly ordered the precipitating action; hecatombs were celebrated with feasting. From that (roughly halfway) point of his “Foreword” on, Simberloff used Elton mainly as a springboard for promoting more recent conceptions. “Elton clearly foresaw some patterns,” Simberloff wrote, “that have since been more comprehensively confirmed and are now the reigning tenets of invasion biology (all are well treated in Williamson’s [1996 book] Biological Invasions)”.637 He mentioned a Mediterranean “killer” alga, but unaccountably failed to promote his then-recent University of Chicago Press translation of a book on the topic, even by reference. Then Simberloff “dropped the big one,” invasion biology’s “mother of all” statements of faith. Promulgated narrowly (and irreproducibly) by his own mentor in 1992, and since extrapolated, modified and asserted willy-nilly, it goes like this: “We now believe that introduced species are the second greatest cause of species extinction and endangerment (following habitat destruction) both in the United States and worldwide.”638 If a conception can be invasive, this one certainly was.639 636 Simberloff, “Foreword” (cit. n. 627) ix. 637 Ibid., x. 638 Ibid., xi. 639 M. K. Chew, in preparation, “The Invasion of the Second Greatest Threat.” 281 “What did Elton get wrong, and what did he leave for us to do other than mop-up operations?” Simberloff asked, identifying his intended audience as fellow heirs of the Eltonian legacy.640 For the present purpose, analyzing a litany of the things Elton did not know, suspect, recognize or understand in 1958 is not terribly important. But it preoccupied Simberloff, whose agenda required Elton’s The Ecology of Invasions only as a point of departure. The list included another Simberloff preoccupation, the “synergistic interactions among invaders [that] may well lead to accelerated impacts on native ecosystems – an invasional meltdown process.” 641 The play on Elton’s own weak framing metaphor of nuclear explosions diminished to nuclear power plants is an entertaining one, and the purposefully alarmist conception was doubtless a welcome addition to Carlton’s Biological Invasions 1(1). “So,” Simberloff concluded, “though Elton sketched the broad outlines of the scientific questions asked, and the nature of the research to answer them, there is still much work for invasion biologists to do.” 642 And later in the paragraph, “much research is needed to make these tools [quarantine, eradication, suppression] effective. Never mind that the only specific calls for future research Elton made in Invasions were into the relationship between complexity (or variety) and stability.643 Never mind that Elton thought he had an 640 Ibid., xi. 641 Simberloff and Von Holle. “Invasional Meltdown?” (cit. n. 325). 642 Simberloff, “Foreword” (cit. n. 627) xi. 643 Elton, Invasions (cit. n. 162),, 146; 153. 282 adequate handle on the problem, and used his last two chapters to impart an elaborate argument for better conservation, not for more or better science. Simberloff closed with what appears to be vague condescension and dissembling, and a final expression of historical illiteracy: “Anyone wishing to understand the problem, from ordinary citizens through specialized researchers, can profit from reading (or rereading) The Ecology of Invasions. It imparts a wealth of timely information, while Elton’s erudition and dry wit are periodically dazzling. A writer who can describe oysters as ‘a kind of sessile sheep’ and characterize advances in quarantine methods by the proposition that ‘no one is likely to get into New Zealand again accompanied by a live red deer’ is more than just a scientist pointing out an important unrecognized problem” [emphasis mine].644 And perhaps Elton’s advice to Rachel Carson, that authors should preface their own works, was more than just an excuse not to preface hers. Reviews of the 2000 Edition As foreshadowed (and perhaps demanded) by Simberloff’s foreword, invasion biology’s true believers celebrated the return of The Ecology of Invasions to ready availability. Like Simberloff they attributed much to Invasions that Elton and others had pored over or worked out previously. They somehow simultaneously recognized and misconstrued its significance. Reviewers for Conservation Biology, Diversity and Distributions, and American Zoologist all 644 Simberloff, “Introduction” (cit. n. 627) xi. 283 declared it to be the definitive beginning of invasion biology. 645 All three seem to have dutifully taken their cues on how to read it from Simberloff’s commentary. Only one of them noticed the book’s “nearly three hundred references from the primary scientific literature of [Elton’s] day.”646 Crediting author number 301 with originating a major field of study is an interesting ploy. All three reviewers reverently attended to the book’s nonscientific language and accessibility to a wide audience. “It proved extremely easy to read,” wrote one, apparently expecting a turgid relic.647 Another called it “highly entertaining,” continued “it is discouraging that, more than 40 years later, our populace does not appear to be much better informed about the conservation issues associated with invasions than they were in Elton’s time,” and finally “Elton’s book, although sophisticated and packed with novel ideas and hypotheses, was also a passionate clarion call to awaken the public to an environmental crisis happening in their backyards.” 648 Perhaps they were inured to the implications of these statements by the commonness of “infotainment” appeals to public sentiment in their own fields. Two reviewers suggested that Elton’s questions about human redistribution of biota still drove the agenda of invasion biology, forty years down 645 James E. Byers, [Review] “The Ecology of Invasions by Animals and Plants” American Zoologist 41(2001)1242; Ingrid M. Parker, “Invasion Ecology: Echoes of Elton in the Twenty-First Century” (cit. n. 4); Scott Henderson “The Foundations of Invasion Biology Revisited” Diversity and Distributions 7 (2001) 107. 646 Henderson, “Foundations.” (cit. n. 645). 647 Byers, “The Ecology” (cit. n. 645). 648 Parker, “Echoes.” (cit. n. 645). 284 the pike.649 The third protested that progress had been made in the interim, but made the same point in a roundabout way: “If Elton were alive today, he would be satisfied by the continuing relevance of his work, but also by the exciting development of the field he inspired.”650 That is certainly a moot question. Elton the ecologist reveled in ecological detail; but with Invasions he pointedly attempted to inspire a general change in our approach to nature conservation. He sought the creation of an ecological technocracy with the power of command. In that sense Elton’s “Millennium” is not yet at hand, nor is it on the horizon. One more review of the reissued Invasions completes the picture. In a special, eight page “book section” of BioScience, Elton’s Invasions appeared among an inventory of some thirty book-length publications (1983-2003, but mostly post-1999) sorted and evaluated by Daniel Simberloff.651 It was a very Eltonian turn. Biannually for over two decades, Charles Elton and his cadre of reviewers produced annotated “Notices of Publications on the Animal Ecology of the British Isles” for The Journal of Animal Ecology, thereby determining which new literature was worth reading (and despite the title, not only in Britain). Simberloff assumed command of the major publications of invasion biology not merely for the “internal” edification of the field itself, but for a much broader potential audience of biologists in general. Thus, in 2004, Simberloff had the 649 Byers, “The Ecology” (cit. n. 645); Henderson, “Foundations.” (cit. n. 645). 650 Parker “Echoes.” (cit. n. 645). 651 An editorial insertion disclosed that, among the reviewed items, Simberloff had translated one, contributed the foreword to another, two chapters to a third, one chapter to a fourth, and “participated in a workshop that designed material” in another. 285 opportunity to both revise and reify his own prior contextualization of “Charles Elton’s 1958 classic, The Ecology of Invasions by Animals and Plants”: This bible of invasion biology, written by a leading ecologist of the last century, is an expansion of a series of popular BBC radio broadcasts. Elton vividly sketches the three basic components of invasion biology: the evolution of distinct biotas in isolation, the shattering of that isolation by human trade and travel, and the disastrous impacts of some of this mixing. Many of Elton’s examples remain timely, and he foresaw some catastrophic subsequent invasions (for instance, his concern with fouling organisms and ballast water was prescient). The generalizations in Elton’s popular book — such as the high invasibility of disturbed areas— are almost point-by-point the same ones invasion biologists propound a half century later. The only major current topic he did not address is the evolution of invaders in their new homes. That he synthesized the major themes in a way that inspired much of modern invasion biology, while referring to H.G.Wells, Buffalo Bill, Walt Whitman, Charlie Chaplin, and Pieter Brueghel the Elder, indicates that this is no dry specialist’s monograph.652 “Popular book” was a key judgment. In rapid succession, Simberloff called Invasions “this bible of invasion biology,” retired Elton to “the last century” and relegated his book to the (usually) scientifically disreputable category of “popular accounts” along with Yvonne Baskin’s A Plague of Rats and Rubbervines (quoted in my introduction), Australian Tim Low’s Feral Future, Jason and Roy Van Driesche’s Nature out of Place, and Kim Todd’s Tinkering with Eden.653 But now Simberloff placed Invasions no better than third among these “recent books that aim to bring the great scope and complexity of biological invasions to a broad readership.” To Simberloff, Feral Future and A Plague were 652 Daniel Simberloff, “A Rising Tide of Species and Literature: A Review of Some Recent Books on Biological Invasions” BioScience 54 (2004) 247-254. 653 Yvonne Baskin, A Plague of Rats and Rubbervines, (cit. n. 6); Tim Low, Feral Future, (Chicago, University of Chicago Press, 2002); Jason Van Driesche and Roy Van Driesche, Nature out of Place, (Washington DC: Island Press, 2000); Kim Todd, Tinkering with Eden (New York: Norton, 2001). 286 the standout efforts. To Simberloff, and (given his assumption of a leading role, to invasion biology in general) Elton was no longer a founder, but merely an “inspiration.” Conclusions Statements published by leading invasion biologists at the turn of the twenty-first century promoted the idea that serious scientific interest in neobiota arose in the 1950s. I countered these statements with a three-point thesis: (1) Scientists began observing and theorizing about neobiota no later than the mid-eighteenth century. (2) Charles Elton “pulled his thoughts together” regarding neobiota episodically, and published them several times, beginning in the 1920s. (3) Invasion biologists and their allies looked no further than Elton and The Ecology of Invasions by Animals and Plants for a “bible” and a founder because the erewhile celebrated ecologist’s alarmist book provided authoritative support for their own alarmist responses to neobiota. To support point one, I provided evidence for over two centuries of scientific investigation and publication regarding neobiota, some (but not all) of which anticipated Elton’s concerns. To support point two, I showed how Elton’s martial and otherwise negative conceptions of neobiota formed very early in his career, and appeared in all of his earlier books and several papers. To support point three, I showed how Daniel Simberloff re-framed The Ecology of Invasions for an audience of invasion biologists in his foreword to the book’s 2000 reissue, and how it was then received by modern reviewers. Many subsidiary questions arose in the course of the project, but the major ones were: What vestiges of pre-Eltonian work persist in modern invasion biology? Why are Elton’s “reasons for conservation” so strongly reminiscent of 288 Aldo Leopold’s “land ethic”? Why did Elton to take up the issue of human redistribution of biota as a book project in the late 1950s? Remnants of pre-Eltonian work survive in the modern practices of invasion biology, but not necessarily because Elton passed them forward; he seems to have been largely unaware of most earlier and even parallel work, especially by botanists. The earliest and most extensive, continuing studies of neobiota were botanical, but although Elton included “plants” in the title of his book, he discussed very few such cases. He based much of his argument on Alfred Russell Wallace’s conception of zoogeographic realms, but did not cite Wallace’s own disquiet over “the fate of remote islands.” A concern with sorting “the biota of places and the places of biota” (to paraphrase H.C. Watson) was initiated in the 1840s, and survives today both in the intuited distinction between alien and native species, and in a more elaborate system of concepts and terms denoting the attachments between biota, generalized habitats and geographical regions. Most such terms are rendered from Greek roots, so I named the practice anekeitaxonomy, from the Greek ανήκει, “belonging.” Scientists’ motivations for doing anekeitaxonomy have evolved. Centers of evolution have replaced centers of creation, but are often regarded as no less metaphysically significant. In the nineteenth century, Wallace (again), and J.D. Hooker broached the scientific value of preserving endemism and biotic uniqueness on oceanic islands. Defending such uniqueness in terms of a “sense of place” approached a moral imperativeness in New Zealand by the 1920s, and 289 that imperative later expanded elsewhere under rubrics including the “land ethic” and more recently, “biodiversity” protection. Elton’s reason’s for conservation actually were Aldo Leopold’s “land ethic,” and vice versa. Enough documentary evidence was found to show that their ideas emerged from a “competitive collaboration,” a relationship to which each also brought an instinctive distaste for neobiota. Although they never coauthored a paper, the two interacted as friends and colleagues, bemoaning neobiota and converging upon a shared conservation philosophy. In the late 1950s Elton was provoked to write again about neobiota when (among other incentives) U.S.-born, neotropical ecologist Marston Bates produced a brief but comprehensive, new and relatively objective analysis of the topic. By any standard, but certainly by comparison, Elton’s resulting Ecology of Invasions was a polemic. Elton and Bates corresponded and interacted personally before and after the fact of their rival publications, and continued to correspond afterward, although I found no specific discussion of their different conclusions. An advanced hypothesis, if not a firm conclusion emerged from comparing their lives and their views: Elton’s English middle-class upbringing and career, spent drilling with the Army, doing “war work” and experiencing rapid modernization led him to a conflict-based conception of neobiota; Marston Bates saw a fundamentally different world while working as a purely civilian researcher in the American tropics during a mostly Eurasian war. Bates ultimately supported, or at least 290 deferred to, Elton’s representation, but only after relocating from his beloved tropics to the domesticated farmscape of southern Michigan. Uses for a Dead Celebrity The creation myth commemorating Elton as invasion biology’s ultimate patriarch, and the drafting and publication of The Ecology of Invasions as its genesis event addressed three needs. The commemoration itself appealed to the authority of a legendary (and conveniently deceased) ecologist, claiming the man as an emblem of the new (would-be) field of endeavor. Charles Elton almost always shunned notoriety in favor of privacy and (avowedly) family life. He successfully avoided establishing a strong public persona, to the extent that his published, pre-1958 comments regarding neobiota and the extent of his contributions to British nature conservation were all but forgotten. As a result, Elton was available for retrospective “rebranding,” inviting would-be successors to claim his name and authority via selective quotation and innuendo. Adding the date and/or title of a specific Eltonian opus rendered any prior efforts irrelevant, and (if credulously accepted) also effectively nonexistent. One needed look no further back than 1958, and certainly no farther afield than a vaguely familiar authority figure from traditional mainstream ecology. More subtly, for the cognoscenti, the date situated Elton’s book between Aldo Leopold’ A Sand County Almanac (1948) and Rachel Carson’s Silent Spring (1962), and situated Elton himself between them as another significant prophet of modern environmentalism. It qualified him not merely to describe ecology’s “is,” but to prescribe ecology’s “ought.” Nevertheless, few if any modern invasion 291 biologists have engaged with or explained Elton’s conservation philosophy, which was the primary object of his Invasions exercise and so interested his early reviewers. In the 1990s, the would-be leaders of invasion biology promoted Charles Elton as a culture hero and his book as a “bible” in their pursuit of a collective identity and authority. Modern invasion biology grew from the same commonsense but theoretically feeble assumption that inspired earlier workers: biotas belong to regions, combined with a more recent reciprocal conviction that regions belonged to biotas. Buttressed by credible and/or merely frightening (but relatively ad hoc) practical justifications, invasion biology aspired to practice biogeographical enforcement, a kind of reactionary land reform, an inverse distributive justice. Only after a new wave of biologists produced authoritative accounts taking conceptions of biogeographical belonging for granted and rendering them again tacit could invasion biology begin the process of retiring Elton’s increasingly anachronistic book. Tennessee ecologist Daniel Simberloff assumed a major role in shepherding this process of adopting and then retiring the Elton myth, and began to emerge as a primary authority in his stead. Simberloff attributed his biodiversity-based concerns about neobiota to religious or religion-like convictions. Implications Invasion biology should discard its Eltonian creationism, engage with recorded history, and thus rediscover itself. What is the “real” character of our 292 fascination with the biota of places? Can we see ourselves in Kalm, or Watson, or Wallace, or Thellung, or Bates? Can we find our questions among those they tried to answer? Do we need biogeographical “just so stories” about nature’s (or Nature’s) “is and ought,” and about our own “can and will”? Is writing them a scientific endeavor, an endeavor of scientists, or a religious declaration? What is the difference, and what difference does it make? If we stand too firmly on the shoulders of Charles Elton, we will see only what he saw. The nostalgic scion of a decomposing, militarist empire, he looked at neobiota amidst the fog and wrack and dread of two world wars, and saw invasions. Though neobiota lacked military coordination, territorial ambition, or even geographical awareness, he saw invasions. Marston Bates looked at the very same phenomena, and saw accidents and emerging symbiosis. Both men were ecologists. But at least in those moments, only one was writing as a scientist. Sciences are distinguishable by their use of scientific methods to answer a limited set of questions. Charles Elton’s personal background provided him with a viewpoint, and his scientifically acquired knowledge of population dynamics and his extensive reading about neobiota, “the penalties of an ecological education,” put a set of problems in view. But not every problem a scientist faces is scientifically approachable, and not every method a scientist employs is a scientific method. Elton knew this, and rested his arguments for conservation, his vision of what the world should be like, on religious, aesthetic, and practical criteria. Those criteria informed his research choices, inspiring him to devote 293 part of his career to the practical problem of preventing animal depredations of stored foodstuffs. Scientific methods alone could not generate the proposition that rats should not thrive in grain silos, but neither were they needed to help farmers or food manufacturers decide whether rats should eat their inventories. Scientific methods could be used to test the relative merits of different strategies for implementing an agreed-upon goal of reducing rat populations, which is exactly how Elton and his BAP colleagues applied themselves during World War Two. Still, in The Ecology of Invasions and other works, Elton made confused and therefore confusing cases for his views about neobiota. He presented the demarcations of “Wallace’s Realms” as imperative boundaries rather than descriptive ones. And though a few moments’ reflection reveals that there are fundamental differences between neobiotic phenomena and military incursions, Elton never revised his literally youthful and naïve conception that neobiota were invaders, and habitually described them with full-blown military rhetoric. It is doubtful whether Elton ever recognized the extent to which these errors encumbered his reasoning. Perhaps he was never compelled to escape the influence of Beatrix Potter’s “unblemished” anthropomorphic fables, or his warfare-fed upbringing. But his resulting judgments had Carrollian qualities that would have troubled him, had he seen them. As an eminent scientist, convinced beyond self-correction and later revered (or at least presented as) above skeptical review, his powers exceeded even Humpty Dumpty’s. Unlike Alice’s ovoid interlocutor, Elton was not limited to merely indulging in the solipsistic 294 pursuit of making words mean whatever he wanted them to. His meanings outlived him to help convince the unwary but well intentioned that one particular sort of nonsense is sensible. To the extent that invasion biology seeks to enforce some past distribution of biota upon the future, that power of persuasion must be encouraging. But even simple human actions generate unforeseen and unintended consequences. There are more (and more complex) human actions occurring every day. It will be difficult in practice to escape Charles Lyell’s proposition: “We may regard the involuntary agency of man as strictly analogous to that of the inferior animals. Like them we unconsciously contribute to extend or limit the geographical range and numbers of certain species, in obedience to general rules in the economy of nature, which are for the most part beyond our control.”654 Charles Elton despaired that the necessary future of efforts to fortify biogeographic borders against invasion would be a costly, permanent, ineffective state of siege. Neobiota are both causes and effects of environmental change, but since they aren’t really invading in any useful sense of the term, we must explore alternative conceptions and corresponding responses. Invasion biology’s responses to neobiota are constrained by its militarist central dogma. Until and unless that dogma is reformed, alternative responses will have to arise elsewhere. 654 Lyell, Principles of Geology (cit. n. 22), 122. REFERENCES Allee, W. C. Alfred E. Emerson, Orlando Park, Thomas Park, and Karl P. Schmidt. 1949. Principles of Animal Ecology. Philadelphia: W.B. Saunders. Andrewartha H.G. and L.C. Birch. 1954. The Distribution and Abundance of Animals. Chicago: University of Chicago Press. Anker, Peder. 2001. Imperial Ecology: Environmental Order in the British Empire, 1895-1945. Cambridge, MA: Harvard University Press. Anonymous. und. Sidney Irving Smith. Yale Peabody Museum: History and Archives, www.peabody.yale.edu/archives/ypmbios/smith_si.html, accessed 15 October 2006. Anonymous. und. Texas Tech University class materials accessed online at www.pssc.ttu.edu/pss1411cd/PLANTID/buroak/buroak.htm. 13 September 2006. Anonymous, und. Tyler Prize Homepage, accessed online 25 September 2006 at http://www.usc.edu/dept/LAS/tylerprize/index.html. Anonymous, und. Tyler Prize Laureates (undated) accessed online 25 September 2006 at http://www.usc.edu/dept/LAS/tylerprize/previous.html. Anonymous, 1931. “Brief Notices,” The Quarterly Review of Biology 6: 223. Anonymous, 1932. “Game System Deplored as a Melting Pot” Journal of Forestry 30: 226-227. 296 Anonymous. 1997. Plate Tectonics: The Rocky History of an Idea. University of California, Berkeley, Museum of Paleontology, accessed online 31 August 2006 at http://www.ucmp.berkeley.edu/geology/techist.html Ascherson Paul, Flora der Provinz Brandenburg, der Altmark und des Herzogthums Magdeburg Berlin: A. Hirschwald, 1864 ———. 1883. (untitled) in Johannes Leunis Synopsis der Pflanzenkunde 3, I. Asmous, Vladimir 1945. Dr. E.V. Wulff—A Victim of War. Chronica Botanica 9: 210-211. Avalon Project at Yale Law School, 1996-2005. Nuremberg Trial Proceedings, Volume 18. www.yale.edu/lawweb/avalon/imt/proc/07-10-46.htm. Accessed online 27 September 2006. Baskin, Yvonne. 2002. A Plague of Rats and Rubbervines: The growing threat of species invasions. Washington D.C.: Island Press. Bates, Marston, and Donald Abbott. 1959. Ifaluk: Portrait of a Coral Island. London: Museum Press. Bates, Marston, and The Editors of Life Magazine. 1964. The Land and Wildlife of South America, Life Nature Library. New York: Time, Inc. Bates, Marston. 1945. Observations on Climate and Seasonal Distribution of Mosquitoes in Eastern Colombia. Journal of Animal Ecology 14: 17-25. ———. 1949. The Natural History of Mosquitoes. New York: Macmillan. ———. 1956. Man as an Agent in the Spread of Organisms. In Man's Role in Changing the Face of the Earth, edited by W. L. Thomas, C. O. Sauer, M. Bates and L. Mumford. Chicago: University of Chicago Press. 297 ———. 1960. The Forest and the Sea. New York: Random House. ———. 1961. Man in Nature. Edited by W. D. McElroy and C. P. Swanson. Foundations of Modern Biology. Englewood Cliffs, NJ: Prentice-Hall. ———. 1970. A Jungle in the House. New York:. ———. Papers. University of Michigan, Bentley Historical Library. Bates, Nancy Bell. 1947. East of the Andes and West of Nowhere. New York: Charles Scribner's Sons. Bean, Michael J., and Melanie J. Rowland. 1997. The Evolution of National Wildlife Law. Third ed. Westport, CT: Praeger. Bentham George. de Candolle’s ‘Geographical Botany’ Edinburgh Review or Critical Journal 104 1856 490-518. ———. 1896. Handbook of the British flora. London: L. Reeve & Co. Binney, Amos. 1851. The Terrestrial Air-Breathing Mollusks of the United States, and the Adjacent Territories of North America. 2 vols. Boston: Charles C. Little and James Brown. Binney, George. 1926. With Seaplane and Sledge in the Arctic. New York: George H. Doran Co. Bookchin, Murray, 1997. The Murray Bookchin Reader, Janet Biehl, Ed. London: Cassell. Boym, Svetlana. 2001. The Future of Nostalgia. New York: Basic Books. Brands, S.J. (compiler) 1989-2005. Systema Naturae 2000. Amsterdam, The Netherlands. [http://sn2000.taxonomy.nl/] 4 January 2006. 298 Briggs, John C. and Christopher J. Humphries. Early Classics. IN Lomolino, Mark V., Dov F. Sax, and James H. Brown, eds. 2004. Foundations of Biogeography: Classic Papers with Commentaries. Chicago: University of Chicago Press. British Ecological Society. 1944. Nature Conservation and Nature Reserves. The Journal of Ecology 32 (1): 45-82. Bull, Peter. 2006. Wodzicki, Kazimierz Antoni z Granowa 1900 - 1987. Dictionary of New Zealand Biography updated 7 April 2006; accessed online 16 October 2006 at http://www.dnzb.govt.nz/. Byers, James E. 2001. Book Review: The Ecology of Invasions by Animals and Plants by Charles Elton. American Zoologist 41:1242. Cadotte, Marc William. 2006. Darwin to Elton: Early Ecology and the Problems of Invasive Species, IN Marc William Cadotte, Sean M. McMahon and Tadashi Fukami, eds. Conceptual Ecology and Invasion Biology: Reciprocal Approaches to Nature Dordrecht: Springer, 15-33. Campbell, Douglas H. 1926. An Outline of Plant Geography. New York: Macmillan. Candolle, Alphonse de. 1855. Geographie Botanique Raisonée Paris: Masson. ———. 1885. Origin of Cultivated Plants. New York: D. Appleton. Candolle, Augustin P. de. 2004. Elements of the Philosophy of Plants, in Mark V Lomolino, Dov F. Sax, and James H. Brown, eds. Foundations of Biogeography Chicago: University of Chicago Press. 299 Carlton, James T. 1999. Editorial: A Journal of Biological Invasions. Biological Invasions 1: 1. Carpenter, Humphrey 1997. The Envy of the World: Fifty Years of the BBC Third Programme and Radio 3 (London: Phoenix Giant. Carrington, John T. 1893. Books to Read. Science-Gossip, n.s. 1:15-16. Carr-Saunders, A.M. 1922. The Population Problem (Oxford: Clarendon). Castro, Josué de. 1952. The Geography of Hunger Boston: Little, Brown, Chew, Matthew K. and Manfred D. Laubichler. 2003. Natural Enemies- Metaphor or Misconception? Science 301:52-53. Chitty, Dennis. 1996. Do Lemmings Commit Suicide? Beautiful Hypotheses and Ugly Facts. Oxford: Oxford University Press. Clark, Andrew Hill. 1949. The Invasion of New Zealand by People, Plants and Animals: The South Island. Westport, CT: Greenwood Press. ———. 1956. The Impact of Exotic Invasion on the Remaining New World MidLatitude Grasslands, in Thomas, et al, Eds., Man’s Role in Changing the Face of the Earth, 737-762. Clements, Frederick. 1916. Plant Succession Washington D.C.: Carnegie Institution. Cohn, Margaret Foreman. 1959. Book Review: “The Ecology of Invasions by Animals and Plants” by Charles S. Elton.” The Quarterly Review of Biology 34: 303-304. 300 Congress of the National Socialist Workers' Party of Germany. 1935. The Reich Citizenship Law of September 15, 1935. Accessed online 5 February 2006 www.mtsu.edu/~baustin/nurmlaw2.html. Cooke, May T. 1928.The Spread of the European Starling in North America United States Department of Agriculture, Circular No. 40, accessed 4 Sept. 2006 at http://www.wku.edu/~smithch/biogeog/COOK1928.htm. Coulter, John M., Henry C. Cowles, and Charles R. Barnes. 1911. A Textbook of Botany for Colleges and Universities. Vol. 2. New York: American Book Company. Cox, David L. 1979. Charles Elton and the Emergence of Modern Ecology. Dissertation, Dept. of Biology, Washington University, Saint Louis, MO. Crowcroft, Peter. 1991. Elton's Ecologists. Chicago: University of Chicago Press. Darwin, Charles. 1859. On the Origin of Species by Means of Natural Selection London: John Murray. ———. 2004. The Voyage of the Beagle. A. Brandt, Ed. Washington DC: National Geographic Society. Davies, J. L. 1958. The Pinnipedia: An Essay in Zoogeography Geographical Review 48: 474-493 Davis, Fred. 1979. Yearning for Yesterday: A Sociology of Nostalgia New York: The Free Press. Davis, Mark A. 2006. Invasion Biology 1958-2004: The Pursuit of Science and Conservation. In Conceptual Ecology and Invasions Biology: Reciprocal 301 Approaches to Nature, edited by M. W. Cadotte, S. M. McMahon and T. Fukami. Dordrecht: Springer. Davis, Mark A., Ken Thompson, and J. Philip Grime. 2001. Charles S. Elton and the dissociation of invasion ecology from the rest of ecology. Diversity and Distributions 7:97-102. Drake James, Foreword IN Marc William Cadotte, Sean M. McMahon and Tadashi Fukami, eds. Conceptual Ecology and Invasion Biology: Reciprocal Approaches to Nature. Dordrecht: Springer. Drake, J. A., Harold A. Mooney, Francesco Di Castri, R. H. Groves, F.J. Kruger, Marcel Rejmanek, and Mark Williamson, eds. 1989. Biological Invasions: A global perspective. Vol. 37, SCOPE. Chichester, UK: Wiley and Sons. Egerton, Frank N. 1976. Ecological Studies and Observations before 1900. Issues and Ideas in America, ed. B.J. Taylor and T.J. White Norman: University of Oklahoma Press. 311-51. ———. 2003. Hewett Cottrell Watson: Victorian plant ecologist and evolutionist. Aldershot: Ashgate. Egler, Frank E. 1942. Indigene Versus Alien in the Development of Arid Hawaiian Vegetation. Ecology 23 1:14-23. ———. 1961. The Nature of Naturalization. Recent Advances in Botany. IX International Botanical Congress. ed. D.L. Bailey.. Montreal: University of Toronto Press, Elton, Charles S. 1925. The Dispersal of Insects to Spitsbergen. Transactions of the Entomological Society of London 73: 289-299. 302 ———. 1927. Animal Ecology. London: Sidgwick & Jackson, Ltd. ———. 1930. Animal Ecology and Evolution. Oxford: Clarendon. ———. 1933. Alien Invaders. The Times, May 6, 1933, 13-14. ———. 1933. Exploring the Animal World 5: Plagues of Animals. The Listener, May 24 1933, 828-830. ———. 1933. The Ecology of Animals. London: Methuen. ———. Ed. 1933. Matamek Conference on Biological Cycles, Labrador, 1931: Abstract of Papers and Discussions. Canadian Labrador: Matamek Factory. ———. 1942. Voles, Mice and Lemmings: Problems in Population Dynamics. Oxford: Clarendon Press. ———. 1943. The Changing Realms of Animal Life. Polish Science and Learning 2:7-11. ———. 1944. The Biological Cost of Modern Transport. Review of Anopheles Gambiae in Brazil 1930-1940 by Soper, Fred L.; Wilson, D. Bruce. The Journal of Animal Ecology 13 1:87-88. ———. 1952. Mammals in New Zealand. Review of Wodzicki, K. Introduced Mammals of New Zealand. The Journal of Animal Ecology 21 1:161-162. ———. 1957. Radio Scripts. British Broadcasting Corporation Written Archives. ———. 1957. Balance and Barrier: 3. The Conservation of Variety. The Listener, April 11, 1957, 596-597, 560. ———. 1958. The Ecology of Invasions by Animals and Plants. London: Methuen and Co. 303 ———. 1966. The Pattern of Animal Communities New York: Wiley and Sons. ———. 2000. The Ecology of Invasions by Animals and Plants. Chicago: University of Chicago Press. ———. Papers. Oxford University, Bodleian Library. ———. Correspondence. 1932-1962. British Broadcasting Corporation Written Archives. Elton, Mrs. Oliver [Letitia]. 1906. The Story of Sir Francis Drake. London: T.C. & E.C. Jack. Fairchild, David, Elizabeth Kay, and Alfred Kay. 1938. The World was My Garden: Travels of a plant explorer. New York: Charles Scribner's Sons. Fairchild, David. 1930. Exploring for Plants. New York: Macmillan & Co. ———. 1944. Garden Islands of the Great East. New York: Charles Scribner's Sons. ———. 1947. The World Grows Round My Door: The story of the Kampong, a home on the edge of the Tropics. New York: Charles Scribner's Sons. Falls, Cyril. 1944. “The Great World War: Fronts — Western and Eastern.” Illustrated London News (8 July 1944) 46. Fernald, M.L. 1926. Campbell’s Outline of Plant Geography. Ecology 7: 510-516. Fitter R.S.R. 1950. Man’s Additions to the British Fauna Discovery 11: 58-62 ———. 1959. The Ark in Our Midst London: Collins. Flader, Susan, and J. Baird Callicott, eds. 1991. The River of the Mother of God and other essays by Aldo Leopold. Madison: University of Wisconsin Press. 304 Fox, Stephen. 1981. John Muir and His Legacy: The American Conservation Movement. Boston: Little, Brown. Gieryn, Thomas. 1995. Boundaries of Science, IN Sheila Jasanoff, et al, eds. Handbook of Science and Technology Studies. (Thousand Oaks, CA: Sage Publications, 1995.) 393-443. Gröning, Gert, and Joachim Wolschke-Bulmahn. 1992. Some Notes on the Mania for Native Plants in Germany. Landscape Journal 11 (2): 116-126. Hansson, Hans G. 1997. Biographical Etymology of Marine Organism Names (BEMON) Tjärnö Marine Biological Laboratory, accessed 13 October 2006 at www.tmbl.gu.se,. Hardin, Garrett. 1960. The Competitive Exclusion Principle. Science 131: 12921297. Hardy, Alister. 1968. Charles Elton's Influence on Ecology. Journal of Animal Ecology 37: 3-8. Harrop, Angus J. 1926.England and New Zealand from Tasman to the Taranaki War. London: Methuen. Hayward, Ida M., and George C. Druce. 1919. The Adventive Flora of Tweedside Arbroath: T. Buncle. Henderson, Scott. 2001. The Foundations of Invasion Biology Revisited Diversity and Distributions 7: 107. Hesse, Richard. 1937. Ecological Animal Geography. Translated by W. C. Allee and K. P. Schmidt. New York: John Wiley & Sons. Hettinger, Ned. 2001. Exotic Species, Naturalisation, and Biological Nativism. 305 Environmental Values 10:193-224. Highfield, Roger. 2006. Cumbria still affected by fire at Windscale nuclear reactor 49 years on. http://www.telegraph.co.uk/ 25 April 2006. Higham, John. 1963. Strangers in the Land: Patterns of American nativism 18601925. New York: Atheneum. Hooker, Joseph D. 1884. A Student’s Flora of the British Islands London: MacMillan. ———. 1867. Insular Floras. Gardeners' Chronicle January 1867, accessed at www.wku.edu/~smithch/biogeog/HOOK1866.htm, 12 October 2006. Humboldt, Alexander von. 2004. Essai sur la Géographie des Plantes,[1805]. Trans. Francesca Kern and Phillipe Janvier. In Foundations of Biogeography: Classic Papers with Commentaries, Ed. M. V. Lomolino, D. F. Sax and J. H. Brown. Chicago: University of Chicago Press. ———. 1850. Views of Nature: or Contemplations on the Sublime Phenomena of Creation. Translated by E. C. Otté and H. G. Bohn. London: Henry G. Bohn. Huntington, Ellsworth. 1931. The Matamek Conference on Biological Cycles, 1931. Science 74: 229-235. Hutchinson, G. Evelyn. 1959. Homage to Santa Rosalia, or Why are there So Many Kinds of Animals? The American Naturalist 93: 145-159. Huxley, Aldous. 1932. Brave New World. New York: Harper & Brothers. Jacks G.V. and R.O. Whyte. 1939. The Rape of the Earth. London: Faber and Faber. 306 Jax, Kurt. 2002. History of Ecology. In Encyclopedia of Life Sciences. London: MacMillan/Nature Publishing Group. Johnson, Richard I. 2004. The Rise and Fall of the Boston Society of Natural History. Northeastern Naturalist 11:81-108. Kalm, Peter. 1972. Travels Into North America. Translated by J. R. Forster. Barre, MA: The Imprint Society. Kerkkonen, Martti. 1959. Peter Kalm's North American Journey: Its Ideological Background and Results, Studia Historica. Helsinki: Finnish Historical Society. Kimler, William C. 1981. Bates, Marston. In Dictionary of Scientific Biography, edited by F. L. Holmes. New York: Charles Scribner's Sons. 307 Koerner, Lisbet. 1994. Linnaeus' Floral Transplants. Representations 47: 144169. Larson, Brendon M. H. 2005. The War of the Roses: Demilitarizing Invasion Biology. Frontiers in Ecology and the Environment 3: 495-500. Larson, Brendon M. H. Brigitte Nerlich, and Patrick Wallis. 2005. Metaphors and Biorisks: The War on Infectious Diseases and Invasive Species. Science Communication 26 3:243-268. Lear, Linda. 1997. Rachel Carson: Witness for Nature New York: Henry Holt. Leclerc Georges Louis, comte de Buffon. 1775-1776. The natural history of animals, vegetables, and minerals; with the theory of the earth in general. Transl. W. Kenrick and J. Murdoch. London: T.Bell. Eighteenth Century Collections Online. Gale Group. Leopold, Aldo. 1918. Mixing Trout in Western Waters. Transactions of the American Fisheries Society 47:101-102. ———. 1933. Game Management. New York: Charles Scribner's Sons. ———. 1935. Book Review: “Exploring the Animal World” by C.S. Elton. IN S. Flader and J. B. Callicott, eds. 1991. The River of the Mother of God and Other Essays by Aldo Leopold. Madison: University of Wisconsin Press. ———. 1966. A Sand County Almanac, with other essays on conservation from Round River. New York: Oxford University Press. ———. Papers. University of Wisconsin Archives. Lever, Christopher. 1987. Naturalized Birds of the World. Harlow, UK: Longman Scientific & Technical. 308 Lever, Christopher. 2003. Naturalized Reptiles and Amphibians of the World. Oxford, UK: Oxford University Press. Levi, Herbert W. 1952. Evaluation of Wildlife Importations. The Scientific Monthly 74 6:315-322. Lindroth Carl H. 1957. The Faunal Connections Between Europe and North America. New York: John Wiley and Sons. Link, Vernon B. 1955. A History of Plague in the United States of America. Public Health Monograph 26 Washington DC: U.S. Department of Health, Education and Welfare. Linsdale, Jean. 1928. "Some Environmental Relations of the Birds in the Missouri River Region." Wilson Bulletin 40: 157-177. Lomolino, Mark V., Dov F. Sax, and James H. Brown, eds. 2004. Foundations of Biogeography: Classic Papers with Commentaries. Chicago: University of Chicago Press. Low, Tim. 2002. Feral Future: The Untold Story of Australia's Exotic Invaders. Chicago: University of Chicago Press. Lutts, Ralph H. 1990. The Nature Fakers: Wildlife, Science & Sentiment. Golden, CO: Fulcrum. Lyell Charles, Principles of Geology. Vol. 2. London: John Murray, 1832 Macdonald I.A.W. F.J. Kruger and A.A. Ferrar. 1986. The Ecology and Management of Biological Invasions in Southern Africa. Capetown: Oxford University Press. Macdonald, Helen. Falcon. London: Reaktion. 309 Martiniussen, Erik. 2003. Sellafield. Oslo: The Bellona Foundation. McIntosh, Robert P. 1985. The Background of Ecology. Cambridge, UK: Cambridge University Press. Meine, Curt. 1988. Aldo Leopold: His life and work. Madison, WI: University of Wisconsin Press. ———. 2002. Aldo Leopold and the Huron Mountain Club (interview). Environmental Review 9 (7): 1-8. Middleton, A.D. 1931. The Grey Squirrel: The Introduction and Spread of the American Grey Squirrel in the British Isles. London: Sidgwick & Jackson. Milton, Kay. 2002. Loving Nature: Towards an Ecology of Emotion. London: Routledge. Monmonier, Mark. 1996. How to Lie with Maps. 2nd ed. Chicago, IL: University of Chicago Press. Morrell, Jack. 1997. Science at Oxford 1914-1939. Oxford: Clarendon. Mühlenbach, Viktor. 1979. Contributions to the Synanthropic Adventive Flora of the Railroads in St. Louis, Missouri, U.S.A. Annals of the Missouri Botanical Garden 66 1: 1-108. 310 Murie, Olaus J. 1951. Tampering with Nature in New Zealand. Review of Wodzicki, K. The Introduced Mammals of New Zealand. Ecology 32 4:767-768. Nicholson E. Max. 1991. Rebellion Against the Ratpack: Obituary of Charles Elton, FRS The Guardian London 13 May 1991. ———. 1957. Review of Thomas, William L. Jr. Editor Man’s Role in Changing the Face of the Earth. The Journal of Ecology 45:955-958. Odum, Eugene P. 1953. Fundamentals of Ecology. Philadelphia: W.B. Saunders. Olin, Dirk and Edwin Black, 2003. “The Origins Of Hate” The American Lawyer 21 (2 September 2003), accessed online 27 September 2006 at www.waragainsttheweak.com/offSiteArchive/law.com/index.htm. Ordway, Samuel H. 1954. Resources and the American Dream New York: Ronald Press. Ordish, George. 1976. The Constant Pest. New York: Scribners. Osborn, Fairfield. 1948. Our Plundered Planet Boston: Little, Brown. ———. 1953. The Limits of the Earth Boston: Little, Brown. Oxford English Dictionary Online. Oxford: University Press. http://dictionary.oed.com. Various access dates 2001-2006. Parker, Ingrid M. 2001. Invasion Ecology: Echoes of Elton in the Twenty-First Century. Conservation Biology 15: 806-807. Pauly, Philip J. 1996. The Beauty and Menace of the Japanese Cherry Trees: Conflicting visions of American ecological independence. Isis 87: 51-73. 311 ———. 2002. Fighting the Hessian Fly: American and British Responses to Insect Invasion 1776-1789. Environmental History 7:378-399. Paviour-Smith, Kitty. 2004. Elton, Charles Sutherland 1900-1991. In Oxford Dictionary of National Biography. Oxford University Press: Accessed 24 November 2004 at www.oxforddnb.com/view/article/49632. ———. 1991. Obituary: Charles Elton. The Independent 13 May 1991. Pearsall, W.H. The Ecology of Invasion: Ecological Stability and Instability New Biology 29 1959, 95-101. Pollard A. F. 1920. A Short History of the Great War. London: Methuen & Co. Porsild, Morten P. 1932. Alien Plants and Apophytes of Greenland. Copenhagen: Reitzels. Pyšek, Petr. 1995. On the Terminology Used in Plant Invasion Studies. In Plant Invasions: General Aspects and Special Problems, edited by P. Pyšek, K. Prach, M. Rejmanek and M. Wade. Amsterdam: SPB Academic. ———. 2003. On the Road to Understanding: From John Wyndham To Reality Diversity and Distributions 9: 251–252. Rejmánek Marcel. 2005. Invasive Plants: What We Know and What We Want to Know American Journal of Botany 92: 901–2. Richards, P.W. 1959. Book Review: Elton, C.S. The Ecology of Invasions by Animals and Plants.. Journal of Ecology 47 2: 525-526. Ridley, Henry N. 1930. The Dispersal of Plants Throughout the World. London: L. Reeve, Ritchie, James. 1920. The Influence of Man on Animal Life in Scotland: A study 312 in faunal evolution. Cambridge, UK: Cambridge University Press. ———. 1931. Birds and Beasts as Farm Pests. Edinburgh: Oliver and Boyd. ———. 1932. The War Upon Wild Animals The Listener 7: 666-667. ———. 1932. Gains and Losses in Animal Life. The Listener 7: 756-757. Roberts, Michael. 1951. The Estate of Man. London: Faber and Faber. Sagoff, Mark. 2002. What's Wrong with Exotic Species? In Philosophical Dimensions of Public Policy, Ed. V. V. Gehring and W. A. Galston. New Brunswick: Transaction Publishers. Salisbury Edward J. 2004. Ridley, Henry Nicholas 1855–1956. Rev. Oxford Dictionary of National Biography. Ed. H. C. G. Matthew and Brian Harrison. Oxford: Oxford University Press. Sargent, Ralph. 1972. Introduction to Kalm, Peter Travels into North America, trans. J.R. Forster Barre, MA: Imprint Society. Sauer, Carl O. Marston Bates, and Lewis Mumford. 1954. An International Symposium: Man's Role in Changing the Face of the Earth. Man 54:139. ———. 1955. Man's Role in Changing the Face of the Earth: An International Symposium. Science 121: 356-7. ———. 1955. Symposium: Man's Role in Changing the Face of the Earth. Nature 175: 754. Schinz, Hans. 1928. Albert Thellung 1881-1928. Vierteljahresschrift der Naturforschenden Gesellschaft in Zürich 73:558-580. 313 Sclater, Philip Lutley. 1858. On the General Geographical Distribution of the Members of the Class Aves. Journal of the Proceedings of the Linnean Society: Zoology 2: 130-145 ———. 1897. On the Distribution of Marine Mammals, Science 5: 741-748. Shelford, Victor. 1913. Animal Communities in Temperate America. Chicago: University of Chicago Press. Simberloff, Daniel. (und.).(unt.). http://eeb.bio.utk.edu/simberloff.asp Accessed 21 October 2006. ———. 1991. Keystone species and community effects of biological introductions. IN Assessing Ecological Risks of Biotechnology, ed. L. R. Ginzburg. Boston: Butterworth-Heinemann. 1-19. ———. 1992. Conservation of pristine habitats and unintended effects of biological control. In Selection Criteria and Ecological Consequences of Importing Natural Enemies, ed. W. C. Kauffman & J. E. Nechols. Lanham, MD: Entomological Society of America pp. 103-117; ———. 2000. Foreword in Charles Elton, The Ecology of Invasions by Animals and Plants Chicago: University of Chicago Press ———. 2000. Introduction to Charles S. Elton, The Ecology of Invasions by Animals and Plants. Chicago: University of Chicago Press, ———. 2004. A Rising Tide of Species and Literature: A Review of Some Recent Books on Biological Invasions BioScience 54: 247-254 Simberloff, Daniel and Betsy Von Holle. 1999. Positive Interactions of Nonindigenous Species: Invasional Meltdown? Biological Invasions 1: 21-32. 314 Simberloff, Daniel and Peter Stiling. 1996. Risks of Species Introduced for Biological Control, Biological Conservation 78: 185-192. Simberloff, Daniel, and Peter Stiling. 1996. How Risky is Biological Control? Ecology 77: 1965-1974. Slobodkin, L.B. 2001. The Good, the Bad, and the Reified. Evolutionary Ecology Research 3:1-13. Smith, Sidney I. 1868. The Geographical Distribution of Animals. American Naturalist 2:124-31. Southwood, Richard and J. R. Clarke. 1999. Charles Sutherland Elton. 29 March 1900-1 May 1991. Biographical Memoirs of Fellows of the Royal Society, 132-140. Speirs, E. Yvonne. 1993. 'Thomson, George Malcolm 1848 - 1933'. Dictionary of New Zealand Biography, updated 7 April 2006; http://www.dnzb.govt.nz/. Spiro, Jonathan P. 2000. "Patrician Racist: The Evolution of Madison Grant." Ph.D. diss., Dept. of History, University of California, Berkeley. Stead, David G. 1935. The Rabbit in Australia. Sydney: Wynn. Steere, William C. 1953. Douglas Houghton Campbell 1858-1953. The Bryologist 56:127-133. Stevens, P. F. 2004. Bentham, George 1800–1884. Oxford Dictionary of National Biography. Ed. H. C. G. Matthew and Brian Harrison. Oxford: Oxford University Press. Subramaniam, Banu. 2001. The Aliens Have Landed! Reflections on the Rhetoric of Biological Invasions. Meridians: 2 :26-40. 315 Sukopp, Herbert. 2002. On the Early History of Urban Ecology in Europe. Preslia 74: 373–393. Takacs, David. 1996. The Idea of Biodiversity: Philosophies of Paradise Baltimore: Johns Hopkins University Press. Tansley, A. G. 1935. The Use and Abuse of Vegetational Concepts and Terms. Ecology 16(3): 284-307. ———. 1946. Our Heritage of Wild Nature. London: Readers Union/Cambridge University Press. Taylor, Archer. 1975. Selected Writings on Proverbs by Archer Taylor, Wolfgang Mieder ed. Helsinki: Suomalainen Tiedeakatemia. Taylor, Walter P. 1959. Book Review: Elton, C.S. The Ecology of Invasions by Animals and Plants.. Ecology 40: 168-169. Thellung, Albert. 1912. La Flore Adventice de Montpellier. Cherbourg: Le Maout. ———. 1918/19. Zur Terminologie der Adventiv– und Ruderalfloristik. Allgemeine Botanische Zeitschrift für Systematik, Floristik, Pflanzengeographie, etc. 24/25: 36-42. ———. 1930. Die Entstehung der Kulturpflanzen Munich: F.P. Datterer. Theodoropoulos, David. 2003. Invasion Biology: Critique of a Pseudoscience. Blythe, CA: Avvar. Thomas, William L. 1956. Introductory: About the Symposium; About the People; About the Theme. IN Man's Role in Changing the Face of the Earth, Ed. W. L. Thomas. Chicago: University of Chicago Press. Thomson, George M. 1921. Wild Life in New Zealand. Part I - Mammalia. 316 Wellington, NZ: New Zealand Board of Science and Art. ———. 1922. The Naturalization of Animals and Plants in New Zealand. Cambridge, UK: Cambridge University Press. ———. 1926. Wild Life in New Zealand. Part II - Introduced Birds and Fishes. Wellington, NZ: New Zealand Board of Science and Art. Titcomb, John W. 1918. Proceedings of the Forty-Seventh Annual Meeting of the American Fisheries Society, St. Paul, Minnesota, August 29, 30 and 31, 1917.” Transactions of the American Fisheries Society 47: 51-79. Todd, Kim. 2001. Tinkering With Eden: A Natural History of Exotics in America. New York: W.W. Norton. University of Wisconsin, Madison. Archives and Records Management. 2006. Aldo Leopold Papers finding aid. http://digital.library.wisc.edu/ 1711.dl/WIArchives.LeopoldPapers. Van Driesche, Jason and Roy Van Driesche. 2000. Nature Out of Place: Biological Invasions in the Global Age. Washington, DC: Island Press. Vogt, William. 1948. Road to Survival New York: William Sloane Associates. Wallace, Alfred Russell. 1859 A Letter from Mr. Wallace Concerning the Geographical Distribution of Birds Ibis 1: 449-454. ———. 1876. The Geographical Distribution of Animals London: MacMillan. ———. 1881. Island Life, New York: Harper. ———. 1889. Darwinism, New York: Macmillan. ———. 1893. Preface, to Harry Wallis Kew, The Dispersal of Shells London: Kegan Paul, Trench, Trübner, v-viii. 317 Wallace, George. 1990. "Law Enforcement and the `Authority of the Resource,'" Legacy: Journal Of The National Association For Interpretation 1(2): 4-8. Warming Eugenius, 1909. Œcology of Plants Trans. Percy Groom and Isaac B. Balfour. Oxford: Clarendon Press. Watson, Hewett C. 1847. Cybele Britannica Vol. I. London: Longman & Co. ———. 1859. Cybele Britannica Vol. IV. London: Longman & Co. ———. 1868. Compendium of the Cybele Britannica. Vol I. Thames Ditton: privately printed. Wells, H. G., Julian S. Huxley, and G.P. Wells. 1934. The Science of Life. New York: Literary Guild. Wilcove, David S. David Rothstein, Jason Dubow, Ali Phillips, and Elizabeth Losos. 1998. Quantifying Threats to Imperiled Species in the United States. BioScience 48 8:607-615. Willdenow, Karl Ludwig. 1805. Principles of Botany trans. anon. Edinburgh: William Blackwood. Wilson, Edward O. 1958. The Fire Ant. Scientific American 198:36-41 ———. 1992. The Diversity of Life. Cambridge: Belknap. Wilson, R.M. 2005. Retrospective Review: Man’s Role in Changing the Face of the Earth Environmental History 10: 564-566. Winston, Mark L. 1997. Nature Wars: People vs. Pests. Cambridge: Harvard University Press. Wodzicki, K.A. 1950. Introduced Mammals of New Zealand. Wellington: Department of Scientific and Industrial Research. 318 Worster, Donald. 1994. Nature’s Economy. Second Edition. Cambridge: Cambridge University Press. Wulff, Evgeniĭ. V. 1943. An Introduction to Historical Plant Geography. Trans. E. Brissenden. Waltham, MA: Chronica Botanica. Yoon, Carol Kaesuk. 1991. Do Earth’s Species Face Death? Oregonian 15 August 1991, E 1-2. Zohary, Michael. 1962. Plant Life of Palestine, Israel and Jordan. New York: Ronald Press.