Darwin's Corals: A New Model of Evolution and the Tradition of Natural History 9783110680317, 9783110643343

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Table of contents :
Table of Contents
Preface to the English Edition
Preface(2005)
I. The Find (1834)
II. From tree to coral (1837)
III. Strickland’s alternatives (1840)
IV. From the circle drawings to the diagram (1851–1858)
V. The fold-out diagram in the Origin of Species (1859)
VI. Coral: Tradition and Encounter
Conclusion: The lure of the “endless”
Acknowledgements
Bibliography
Index
Picture Credits
Recommend Papers

Darwin's Corals: A New Model of Evolution and the Tradition of Natural History
 9783110680317, 9783110643343

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Horst Bredekamp

Darwin’s cor als

Horst Bredekamp

Darwin’s cor als A new model of evolution and the tradition of Natural History Translated, edited, and adapted by Elizabeth Clegg

This publication was made possible by the Image Knowledge Gestaltung. An Interdisciplinary Laboratory Cluster of Excellence at HumboldtUniversität zu Berlin (sponsor number EXC 1027/1) with financial support from the German Research Foundation as a part of the Excellence Initiative.

ISBN 978-3-11-064334-3 Library of Congress Control Number: 2019946359 Bibliografische Information der Deutschen Nationalbibliothek The Deutsche Nationalbibliothek lists this publication in the Deutsche Nationalbibliografie; detailed bibliographic data are available on the Internet at http://dnb.dnb.de. © 2019 Walter de Gruyter GmbH, Berlin/Boston Deutsche Erstausgabe: Verlag Klaus Wagenbach 2005 Cover illustrations: Melitodes flabellifera var. cylindrica (s. Fig. 11); Charles Darwin. Third sketch of evolution (s. Fig. 14); Charles Darwin, ca. 1881, photography by Elliot & Fry. Design: Petra Florath, Stralsund Printing and binding: Beltz Grafische Betriebe GmbH, Bad Langensalza www.degruyter.com

Table of Contents Preface to the English Edition

vii

Preface (2005)

ix

I. The find (1834)

1

II. From tree to coral (1837)

7

1.  The tree model 2.  Darwin’s first sketches 3.  The model based on coral

III. Strickland’s alternatives (1840) 1. A critique of metaphor 2.  From tree to map 3.  The cartographic model

IV. From the circle drawings to the diagram (1851–1858) 1.  The circle segments 2.  Darwin re-reads Wallace 3.  The precision of Darwin’s initial diagram

V. The fold-out diagram in the Origin of Speci es (1859) 1.  Form 2. Relation to the text 3. A “coral model” for evolution

7 16 18 31 31 34 35

39 39 46 52

61 61 66 68

VI. Coral: Tradition and Encounter 1.  The art of transformation 2.  The cult of coral and the vogue for aquaria 3.  Darwin’s own image of coral as an artist

73 73 77 81

Conclusion: The lure of the “endless”

87

Acknowledgements

91

Bibliography Index Picture credits

93 111 118

PREFACE TO THE ENGLISH EDITION The aim of Darwin’s Corals, which appeared in its original, German edition in 20051, is to investigate Charles Darwin’s preference for a model of evolution (or, in the terminology of his own era, “species transmutation”) that is inspired by the structure and the growth pattern of coral, rather than by those of a tree. Its wider concern is to rescue, for the life sciences, a visual metaphor that stands in contradiction to the attempt to comprehend and represent the natural world in a sociopolitically inflected hierarchical fashion. The continuing need for such an effort to argue the case for Darwin’s “coral model” was recently demonstrated through the evidence of how persistent the allure of the “tree model” has, nonetheless, proved to be. Towards the start of David Quammen’s volume of 2018, The Tangled Tree, this author mentions the “coral model” only to conclude that Darwin very soon discarded this in favour of a “tree model”: “And then he let the coral idea slide, abandoning that metaphor. / The tree of life [metaphor] was better”.2 This statement is more specifically related to Darwin’s celebrated notebook diagram of 1837 headed with the words “I think” (see Fig. 14 of the present volume). But there can in fact be little doubt as to the far from arboreal character of the branching structure committed to that page (nor do Darwin’s related notes make any such claim), while it is not at all difficult to see in it a coralline form. In short: the diagram that Quammen sees as his “crown witness” to Darwin’s switch to a “tree model” is in fact far closer – in its structure and in what we may reasonably surmise of its growth pattern – to a model based on coral. Encouragement for supporters of this last has, however, emerged in new thinking, among scientists, on the question of which evolutionary model or metaphor appears the most apposite in the light of the latest findings in relevant fields. In 2007 the notional “tree of life” was castigated as the emblem of a prolonged misun1   Darwins Korallen. Die frühen Evolutionsdiagramme und die Tradition der Naturgeschichte, Berlin 2005 (2nd edn. 2006). 2   David Quammen, The Tangled Tree. A Radical New History of Life, New York and London 2018, p. 8.

viii

PREFACE TO THE ENGLISH EDITION

derstanding because it fails to accommodate the primarily horizontal movement in genetic exchange between micro-organisms.3 In 2008 an examination of gene transfer among mono-cellular entities concluded that the “tree paradigm fits only a small minority of the genome at best”.4 And in 2009 a further discussion of horizontal gene transfer expressly invoked Darwin’s own reference to “the coral of life”.5 One could, indeed, cite further instances in which similar conclusions have been drawn. The role of coral as a model for all of nature has now assumed a far graver resonance than it had even just fifteen years ago, when I was completing the last of my Darwin research. There can be few parts of our planet where there is not already some awareness, through frequent reports in the press and media, of how the world’s coral formations are suffering through the repercussions of environmental pollution and global warming, almost all of it the reckless contribution of that long-supposed pinnacle of Creation: humankind. As will be clear to any anglophone reader who is able, to some extent, to access texts in German, the present book is a much enriched adaptation of that of 2005 that was its starting point. For this happy outcome I am indebted to Elizabeth Clegg, who brought to the devising of an English version not only her linguistic skills and her scholarly acumen, but also a very real intellectual delight in the subject as I had approached it. I have every hope that her enthusiasm will in due course be shared by many others. Berlin, June 2019

  Ford Doolittle and Eric Bapteste, “Pattern pluralism and the Tree of Life hypothesis”, Proceedings of the National Academy of Sciences of the United States of America, CIV / 7 (2007), pp. 2043–49. 4   Tal Dagan, Yael Artzy-Randrup and William Martin, “Modular networks and cumulative impact of lateral transfer in prokaryote genome evolution”, Proceedings of the Academy of Sciences of the United States of America, CV / 29 (2008), pp. 10039–44, here p. 10043. 5   Gregory P. Fournier, Jinling Huang and J. Peter Gogarten, “Horizontal gene transfer from extinct and extant lineages: biological innovation and the coral of life”, Philosophical Transactions of the Royal Society, series B, CCCLXIV (2009), pp. 2229–39. 3

Preface (2005) “Full fathom five thy father lies, Of his bones are coral made, Those are pearls that were his eyes, Nothing of him that doth fade, But doth suffer a sea-change Into something rich and strange [. . .]” Ariel’s song to Ferdinand, The Tempest, Act I, scene 2 Now and again, in any life, there are moments that provoke an unexpected transformation in one’s entire way of thinking. Such was my experience during the winter semester of 2001 / 2002 at Humboldt University in Berlin, where I was leading a seminar on scientific illustrations. One participant happened to remark that among Charles Darwin’s first diagrams of evolution were drawings that appeared to evoke the structure of coral. For me, this casually imparted information was little short of electrifying. In my study of 1993, published two years later in English translation as The Lure of Antiquity and the Cult of the Machine, I had argued that the frequent presence of striking samples of coral among the valued collectibles assembled in many a sixteenth- and seventeenth-century Kunst- und Wunderkammer may itself have encouraged an apprehension of the evolutionary character of the natural world.1 If Darwin, in an attempt to render in visual form his thinking on evolution (which he addressed as the phenomenon of “species transmutation”), had indeed hit upon the model supplied by the structure of coral, then he would also have been rescuing a crucial element in the traditional understanding of Natural History. Even in putting an end to that tradition, he was achieving its consummation.   Bredekamp 1993, pp. 72–76; Bredekamp 1995, pp. 76–80 Cf. Bredekamp 2000.

1

x

Preface (2005)

Just as evolutionary processes in nature had been surmised long before the advent of Darwin, so did Darwin – or so it now began to seem – elect to operate well within the tradition of Natural History. This consideration was the starting point for my enquiry into Darwin’s corals.2

2  Among the publications from which I have drawn particular inspiration are Dolf Sternberger’s exhilaratingly wide-ranging account of the world in the nineteenth century (Sternberger 1938); Dov Ospovat’s pioneering endeavour to locate Darwin in relation to the intellectual history of his age (Ospovat 1981); and Howard Gruber’s profound analysis of the metaphorical aspects of Darwin’s theory (Gruber 1974; Gruber 1988). Two of the chapters in this volume were published, in a different form: one among the papers of the XIX. Deutscher Kongress für Philosophie des Jah­ res 2002 (Bredekamp 2004 Evolutionsdiagramm), and one among those of the Dresdner Kongress über die Semantik der “Dinge” (Bredekamp 2005 Korallen). My work on Darwin was also reflected in my contribution to a Festschrift presented to Hartmut Böhme (Bredekamp 2004 Üppigkeit). The essence of my argument in these pages formed the basis of the inaugural lecture given, in November 2003, to mark my election as a Permanent Fellow of the Wissenschaftskolleg zu Berlin.

I.  The Find (1834) Among the specimens collected by Charles Darwin during his 1831–36 circumnavigation of the globe, as a naturalist informally attached to the British Admiralty’s hydrographic surveying expedition aboard H. M. S. Beagle under the command of Robert FitzRoy, is an organic form characterised by the repeated bi- and trifurcation of its central shaft into shoots that themselves likewise divide, each advancing by way of a sequence of distinct “bosses” (in botanical terms, “conceptacles”) – hence the later Latin appellation (Fig. 1). The overall form of this entity is striking on account of the vigour still evinced by the record of its alternating left / right progress. The chief, leftward sprig itself emits bifurcating arcs, while its apparently stronger rightward counterpart makes a vigorous lunge further to the right, while both extending and dividing almost vertically. Closer to the point where the main shaft has been severed another rightward shoot strikes out horizontally. And, in the angle formed by the meeting of the two main limbs, there appears yet another (albeit now detached), with its own pattern of sequential bifurcation. Even while entranced by the elegance of this specimen, one is drawn to the record of the stages through which it has passed in the process of becoming part, as it now is, of a museum collection. The matte paper (yellowed with age, and doubtless once much paler) throws into relief the reddish colouring of the prize affixed to it. Identified, at lower left, as “Port Desire, Patagonia”, is the place where this item was found; while the inscription “1770. C. Darwin” records the number allotted to the find when formally registered, and the finder’s name. As revealed in Darwin’s later notes on the items collected during the Beagle voyage, he had made this part­ icular discovery in January 1834 on the Atlantic coast of the southern extremity of South America, near the small settlement known to the few British travellers who reached it as Port Desire (now Puerto Deseado, Argentina).1 At the lower right is the designation “Amphiroa Orbignyana”, a name acknow­ ledging Darwin’s French contemporary, the naturalist and explorer Alcide d’Orbigny; 1  See Darwin: “Coralline Algae Notes” (ed. 1987), p. 194. Port Desire had been so named in 1586 by a much earlier British voyager, the privateer Thomas Cavendish, after his own ship.

2

I.  The Find (1834)

and, below this, the abbreviation “Dne”, itself a reference to the French biologist and taxonomist Joseph Decaisne. The inscriptions in their entirety derive, however, not from Darwin himself, but from the hand of his Irish contemporary and close friend William Henry Harvey.2 As Keeper of the Herbarium at Trinity College Dublin, to which Darwin eventually donated some of his collection and his notes upon it, Harvey was in due course to be responsible for registering the “Corallinae” (including the item in question) among Darwin’s finds.3 Finally, the circular “Herbarium Hookerianum” stamp, with its date of 1867, signals that Darwin’s specimen was one of the duplicates among his finds to be subsequently donated to the magnificent collection formed by William Hooker, who in 1841 was appointed Director of the Royal Botanic Gardens, Kew, on the south-western fringes of London. This collection was formally absorbed by that institution in 1867, by which date Hooker (who had died two years earlier) had been succeeded in his post by his talented botanist son, Joseph, who had become one of Darwin’s closest friends. During the 1960s the Darwin holdings were in turn placed on long-term loan to the Natural History Museum,4 which had been opened to the public in 1881, a year before Darwin’s death, in London’s already well-established South Kensington “museum quarter”. In current taxonomic convention, the entity of which Darwin, in 1834, found an example is known as Bossea (or Bossiella) orbigniana, and is classified among the coralline algae: that is to say algae that, like coral, eventually petrify through a process of calcification. There was, in fact, prolonged disagreement as to whether this organism was correctly categorised among corals (and hence animals) or among algae (and hence plants).5 In the mid-eighteenth century, classified as one of the “Corallinae”, it was seen to belong to the animal kingdom, as was coral itself, in respect of the polypifer, or colony of polyps (minute creatures whose bodies, after death, in due course calcify, thereby “constructing” the coral formation). Around 1840 views on this matter began to change.6 Darwin was never entirely certain as to how the “Corallinae” ought to be categorised; and he was, indeed, fascinated by their indeterminate status.7 But he in any case believed his own find, which Decaisne 2 

I am indebted to Jennifer Bryant und Linda M. Irvine for this information. Harvey 1847, pp. VII–VIII; Darwin: “Coralline Algae Notes” (ed. 1987), p. 188; Ducker 1988, p. 14. 4  I am indebted to Jennifer Bryant for this information. 5  Silva 1957, p. 41. For a summary of the evolving nomenclature of this organism, see www. algaebase.org, where it is listed as Bossiella orbigniana. 6  In 1837 the German naturalist Rudolf Philippi (Philippi 1837) had firmly put the case against classifying the “Corallinae” as part of the animal kingdom. 7  In his own notes on the items he had collected during the Beagle voyage, Darwin was reluctant to see the non-calcified segments of the “Corallinae” he had found in March 1833 in the Falkland Islands as polyps, asking almost indignantly: “What claims have they [the “Coralinae”] to be considered as animals?” See Darwin: “Coralline Algae Notes” (ed. 1987), p. 193. In a letter 3 

I.  The Find (1834)

1 _ Amphiroa Orbignyana (later known as Bossea / Bossiella orbigniana). Found by Darwin in January 1834 on the coast of Patagonia. London, Natural History Museum, Darwin 1770

would name as one of the algae, to be among the small corals.8 It will here, therefore, be regarded as the coral that Darwin himself believed it to be. For it is within this early-nineteenth-century instance of (mis-)perception that the cultural-historical significance of Darwin’s find may be seen to lie. In combination, the appearance of this specimen and the traces of the history of its collection account for that almost magnetic attraction exerted by any item in

of 24 July 1834, to his former Cambridge professor, and continuing mentor, John Henslow, in: Darwin: Correspondence, Vol. I (1985), no. 251, pp. 397–403, here 399, 400, Darwin spoke of the ambiguity of the “Corallinae”: “I feel pretty well convinced [that,] if they are not / Plants, they are not [= nor are they] Zoophites [= zoophytes, i.e. plant-animal hybrids as understood at that period]”. 8  In 1842 Decaisne, in his history of research into the “Corallinae”, had argued for the incorporation into this genus of numerous species of algae. It was in this context that he had named the item in question here Amphiroa Orbignyana (Decaisne 1842, no. 14, p. 124). In Harvey’s publication on the algae of the Pacific Ocean, Darwin’s specimen (albeit found on an Atlantic shore) is thus treated as an example of this category, and hence no longer as an animal, but as a plant (Harvey 1847, p. 100, no. 22; plate XXXVIII, illustrating the specimen then in the Herbarium at Trinity College Dublin).

3

4

I.  The Find (1834)

a museum of Natural History. As soon as it has entered such a collection, a naturally occurring object assumes that quality – attributed in Ariel’s song to Ferdinand in Act I, scene 2 of William Shakespeare’s play The Tempest to a drowned, then symbolically transforming human body – of precious, albeit alien, rigidity.9 Anyone researching in such a museum’s storage depots soon comes to experience much the same sensations as were, and still are, to be felt among collections of reliquaries, or when confronted with that combination of man-made and natural objects to be found in every Kunst- und Wunderkammer, or even among the bewilderingly wide range of items to be discovered in many a Print Room. This power of attraction contradicts those theories that dismiss the supposed “aura” of such an entity as no more than the questionable product of human fancy. But one may, nonetheless, go even further and claim that entities such as Darwin’s specimen, in the form in which it is now encountered, may be said to have assumed the character of a “work of art”. Notwithstanding the chronological and contextual distance from England in the time of Darwin, it would not be altogether inappropriate to invoke here the arguments of Leon Battista Alberti, who maintained that, as soon as a naturally occurring object had undergone some such form of deliberate human intervention – and this might be as relatively little as the scratching of lines upon its surface – it can justifiably be seen to occupy a distinct zone located between the natural and the artificial, to qualify (albeit minimally) as a “work of art”.10 In the case of Darwin’s specimen the intervention consists in the fact of his having first sighted, then intently examined, then decisively plucked and removed the organism in question, in order to have it shipped in due course back to England, where it was to embark on the aforementioned stages of its existence as part of a series of collections. Through the sequence of formal acquisition, registration, codi­ fication and preservation, a distinct “aura” may be found to accrue to such an entity, an “aura” that grows more intense with each stage of processing, bringing the item gradually closer to what, in Alberti’s definition, constitutes a “work of art”. As the case of Darwin demonstrates, this effective transformation was by no means a matter of chance. His case offers, indeed, a paradigmatic instance in which to determine that “surplus” (as understood in Gestalt psychology) that attaches to things when they are removed from their original context. As will be argued in what follows, it is barely possible to explore Darwin’s own motivation if one does not grasp to what extent he must also be viewed as a constructive iconologist. In order to test this hypothesis, it was necessary to examine Darwin’s sketches and drawings, something that is often possible only with the aid of a magnifying glass. Such an approach might at first seem surprising in view of the far from captivating quality of the images in question. Of significance for what follows, how  9  10 

Cf. epigraph on page ix. Alberti, De Statua (ed. 1999), pp. 23 / 24.

I.  The Find (1834)

ever, is a firm belief that the value of scientific visualisation is often determined less by a draughtsman’s technical skill than by the capacity of the image produced to spur the thought process.11 As the motoricity of thought is, in general, apparent in the gestures of the drawing hand more swiftly and directly than it is in speech, what may strike one as no more than the slightest detail can often prove to be an essential characteristic. It is this art-historical approach to the ostensibly insignificant and the apparently unintentional – an approach reflecting twentieth-century psychology but finding its most compelling parallel in the nineteenth-century investigation of crime –12 that informs the present essay. Acknowledging the historical dimension of the natural object, this approach may also be understood as a respectful nod towards the morphological precision that, throughout the 1800s, was a characteristic shared by Art History and Biology.

11 

Cf. on this point: Bredekamp 2005 Denkende Hände; Bredekamp 2005 Zeichnende Denk-

kraft. 12 

Ginzburg 1979.

5

II. From tree to cor al (1837)

1. The tree model It was not until Darwin had returned to England from the H.M.S. Beagle expedition (in October 1836), and had then started attempting to arrange and classify the items he had collected during the voyage, that he began to glimpse, and then to formulate, his first notions of what might be the processes involved in species transmutation – what is now loosely termed “evolution” – heretofore understood in a manner only intermittently and speculatively resembling that which Darwin’s work would at length bequeath to posterity. An observation inscribed, in the summer of 1837, in what he had designated his Notebook B, signals Darwin’s rejection of the theory of “willed” transmutation (“le transformisme”) proposed in the early nineteenth century, by the French botanist, zoologist and taxonomist Jean Baptiste Lamarck. Here, Darwin asserted: “changes not result of will of animal, but [of] law of adaptation”.1 Lamarck, as Keeper of the Muséum d’Histoire Naturelle in Paris, had become convinced that animals of every sort (from the most primitive to the most complex) were capable of transmutation in the process of adapting to an environment that was itself continually altering. He had, however, avoided contradicting the venerable account of Creation supplied in the biblical Book of Genesis through interpreting this process not as a gradual emergence of new animal forms, but as a series of changes more or less expressly undertaken by each category of animal as it sought to adapt itself ever better to that place within the natural order to which it had been allotted. By this means Lamarck had succeeded in establishing a notion of the capacity of animals to alter, without thereby being constrained to

1  Cambridge University Library, DAR 121: Notebook B, fol.  21r. Cf. Gruber 1974, p. 142. On later pages of this notebook Darwin observes “my theory [is] very distinct from Lamarcks” (fol. 214), and “With respect to how species are [,] Lamarck’s willing doctrine [is] absurd” (fol.  216). Cf. on this issue in general: Desmond and Moore 1991, pp. 48–60.

8

II. From tree to coral (1837)

2 _ The Caroligninan Ancestors. Miniature in Ekkehard von Aura, Chronica, late twelfth century. Erlangen, Universitätsbibliothek, Ms. 406, fol.  204v

1. The tree model

3 _ Tree of Porphyry. Wood-cut illustration in 1498 Venetian edition of Aquinas, Opuscula

9

10

II. From tree to coral (1837)

sacrifice the Church’s teaching on the Divine Plan for a Creation perfect, from the start, in every particular.2 In the Notebook B observations that appear to follow directly after the first of the aforementioned aspersions on the Frenchman, Darwin explained his objection to the latter’s theory through invoking the image of a vigorously tentacular tree: “[…] organized beings represent a tree. irregularly branched [,] some branches far more branched. – Hence Genera.”3 Through this model, Darwin affirmed his belief that transmutation on a large scale occurred not through developments taking place, in parallel, among numerous individual organic forms, but through deadly competition between one and another – the competition itself eventually issuing (as the end of the cited note succinctly reveals) in the emergence of diverse new genera. It was through his own objections to Lamarck that Darwin was able to feel his way towards envisaging the interconnections between organic entities after the visual model of a tree. In doing so, he was in fact also employing a model that had long been found in numerous fields of scholarship, from the sociological to the biological (to use terms these would only later acquire). A visual tradition dating back to the early Mediaeval period took its bearings from the belief in the Root of Jesse and the Christian “tree of life”. Over the following centuries this gave rise to diverse ways of employing genealogical, or family, trees (Fig. 2); and, from the sixteenth century onwards, these were being co-opted in demonstration of a single, privileged line of succession.4 Of even more significance was to be the tree model proposed by the thirdcentury Neo-Platonist Porphyry, in which every component within the natural world and every concept related to it was shown to occupy an allotted place.5 From its roots in individual living forms, as shown in the wood-cut featured in a 1498 printed edition of the Opuscula of Thomas Aquinas (Fig. 3), the “trunk” of such a tree rose, in turn, through the further categories of “man”, “animal”, “living creature” and “body”, until it attained the crown of “substance”.6 In the terms “Genus” and “Species”, occurring respectively at the upper and the lower left, this rendering of the arbor porphyriana already supplies notions of larger and smaller categories

2 

Junker and Hossfeld 2001, pp. 49–51. Cambridge University Library, DAR 121: Notebook B, fol.  21r. 4  On the tree of life: Goetz 1965, Fig. 117. On the Mediaeval period: Schadt 1982; on the Early Italian Renaissance: Klapisch-Zuber 1991; fundamental on the Early Modern Period: Heck 2000, who emphasises the distinction between the family tree and the gallery of ancestors; for important examples from the House of Habsburg: Barta 2001, pp. 52–57; in general on this matter: Klapisch-Zuber 2004. 5  Zimmermann 1953, pp. 77–78; Urbani 1991; Wöhler 1992, pp. 3–20; Wendler 2003, pp. 4–31. 6  Aquinas 1498, p. 106v; De abore 1991, p. 337. 3 

1. The tree model

that still hold substantial meaning for us today. As the broad generalisation of the Porphyrian model can easily be perceived as presenting a regularly ascending straight line, it was also possible to approach this notion in the form of a ladder or a chain.7 However, as laterally positioned differentiations occurred between the “steps” of such a linea directa, the impression of a tree endured. And, as these “branches” themselves pointed both upwards and downwards, the true origin of such a tree might, depending on one’s point of view, be found at either its upper or its lower “end”. The category of “man” might, accordingly, be viewed as the start of a process of ever broadening generalisation, or as the crowning glory of a process of increasing specification. For centuries the tree model derived from Porphyry thus allowed its exponents to overlook a two-fold problem central to Natural History: that of the apparent earlier existence of animal forms now extinct, and the evidence of those newly emerg­ i­ng. These phenomena would appear to contradict the implications of the biblical narrative, according to which all forms of animal life had been embraced within the original Divine Act of Creation. It was, however, conceived as possible, for the verticality of the “tree”, the “ladder”, or the “chain” to accommodate newly emerging forms of animal life in as far as these might be incorporated, by way of lateral branches, as ever more subsidiary limbs. It was above all the sixteenth-century redis­ ­covery of the dynamic circular and combinatory arboreal model that had been pro­­ posed by the Mediaeval Catalan logician and philosopher Raymond Lully (Ramon Llull) that was to ensure the repeated relevance of visual metaphors. The usefulness of this model lay in its capacity both to create hierarchical conceptual arrangements and to permit a certain elasticity in modes of branching.8 As early as 1764–65 the Franco-Swiss naturalist Charles Bonnet, in his twovolume Contemplation de la Nature, proposed the concept of a “scale of nature”, illustrated in a later edition as an “idea of a scale of natural beings”: “idée d’une échelle des êtres naturels” (Fig. 4). With reference to the “insects” and “shell-fish” that occur near the mid-point of this “scale”, he posed the question: “Does the scale of nature branch as it advances? Might insects and shell-fish be two lateral and parallel branches of this great stem?”9 In 1787 the spatial layering in Bonnet’s “scale”

7 

Lovejoy 1936, passim. Berns 2000, pp. 160–62, 174–75; Weigel 2003, esp. p. 239. Particularly notable (see Siegel 2004, pp. 41, 52) is the nature of the reception of Lully’s ideas with regard to attempts to hold the arboreal motif responsible for every conceptual error detected by later philosophers (Deleuze and Guattari 1980). The wittily titled 2005 exhibition “einfach Komplex” (meaning “simply complex”) ironically imputed to “depicted trees and the depiction of trees (“Bildbäumen und Baumbilden”) precisely that quality that Deleuze and Guattari had associated with the rhizome (“la rhizome”), in which they saw the counter-image of the arboreal model. 9  ”L’Echelle de la Nature se manifesteroit-elle en s’élevant? / Les Insectes & les Coquillages s’eroient-ils deux Branches latérales & paralleles de ce grand tronc?” Contemplation de la Nature, 8 

11

12

II. From tree to coral (1837)

inspired in the Russian naturalist Peter Simon Pallas the momentous notion of a symbolic double-trunked tree, intended as a model of the organisation of living beings and embracing both the plant and the animal kingdoms: “Among all other visual representations of the system of organic bodies, it would probably be best if one envisaged a tree that, from the root up, has a dual trunk, composed of the very simplest plants and animals, and thus itself both animal and vegetal in character, and yet distinct from each of these”.10 In contrast to the new shoots that, in a tree in the natural world, mediate between two of the main branches, the symbolic tree has cross-ribs that do not emerge as branches of either of the two trunks, but merely stand, autonomously, between these: “The trunk, composed exclusively of growths of the first rank, and comprising generations standing close to each other, would also symbolise those generations that are likewise related to it through lateral connection, but have not become interlocked with each other, even though they have, here and there, extended small subsidiary branches”.11 The powerful arboreal model devised by Pallas enabled practitioners of both Botany and Zoology to evolve patterns of organisation that most suited their own disciplines, as was to be demonstrated, also in 1764, by the German polymath Johann Philipp Rüling with his diagram of differentiated forms of branching, which was intended to be read from the top down.12 Yet again in 1764, the French botanist Antoine Nicolas Duchesne presented a “genealogical tree” (“arbre généalo­ gique”) for a single type of plant – the strawberry – using his accompanying commentary to identify characteristics shared by different varieties, as well as those that counted towards their divergence, and noting the role played by the influence of climate in the emergence of variation.13 Amsterdam 1764, Partie III, chap. 20, p. 59; here cited (with minor adaptations) from anon. Eng. trans.: The Contemplation of Nature, London 1766, Vol, I, part III, chap, 20, p. 50. 10  “Pallas 1787, p. 47: “At omnium optime Arboris imagine adumbraretur Corporum organicorum Systema, quae a radice statim, e simplicissimis plantis atques animalibus duplicem, varie contiguum proferat truncum, Animalem & Vegetabilem” (Pallas 1766, p. 23). Cf. Métraux 2001, p. 12. 11  Pallas 1787, p. 48; “Truncus e principallori generum affinium serie confertus, passim pro ramulis exsereret genera, quae istis laterali affinitati juncta interseri tamen non possunt” (Pallas 1766, p. 24). Cf. Métraux 2001, p. 12. 12  “Tabula Phytographica Universalis affinitates ordum naturalium plantarum exhibens”, in: Rüling 1766, after p. 36; cf. Wendler 2003, p. 39. 13  “Si le Capiton VII n’est pas une plante originairement distincte du Fraisier, il est à croire qu’il est né du Fraisier verd, plutôt que de tout autre, ainsi qu’on le voit dans l’arbre généalogique: il a de commun avec lui, la solidité de ses fruits & la force de ses étamines, mais il en differé [= diffère] par sa grandeur, par l’absence des appendices des feuilles que se voient dans l’autre, par la forme des bouquets de fleurs, la disposition des calices à se renverser, & plus encore par la séparation de sexes sur deux individus. / Le Fraisier écarlate X, qui est le Fraisier commun du Canada & de la Virginie, différé [ = diffère], comme on l’a vû, du Fraisier de bois par un grand nombre de petits caractères; cependant, il ne peut guère avoir pris naissance que de cette race, ainsi que je le

1. The tree model

In the longer term, however, the metaphor devised by Pallas proved to have been a step that would incur numerous conceptual problems. For he had, on one hand, effectively insinuated his model of nature into the intellectual tradition of the metaphorical “tree of life”, which was a staple feature of religious thought and which, in the understanding of Christianity, conceived of new life emerging from the dead wood of the Cross.14 On the other hand, his argument came perilously close to the symbolism of growth, as readily embraced in eighteenth-century discourse on advances in manufacture or the steady increase in human knowledge and understanding, even though this was incompatible with the spirit of the Old Testament. By contrast with the circular and combinatory use of the traditional image of the tree, that devised by Pallas was dangerously modern in that, with its notion of a tree with two trunks, it was likely to encourage the idea of growth entailing a degree of transformation. Once again, there emerged a contradiction between the image invoked and its textual explanation regarding what was permissible. Pallas, accordingly, sought to limit the very speculation that his visual model might well have encouraged, emphasising that his “tree” was to be understood as acknowledging and approving the notion of harmonious and stable relationships, not of those undergoing diverse forms of development. However, while it might well, at this date, have been claimed of every model – be its form that of a ladder, or a scale, or a tree – that it served to describe a natural state and not a natural process, the image in every case, and in particular that evoked by Pallas, nonetheless offered more than was explicitly permitted in the accompanying verbal exposition. Lamarck, who in 1794 collaborated on a French translation of Pallas’s account of his travels as a naturalist, was the first to succeed in closing this gap; and, in doing so, he was able to develop not only a concise theory of transmutation in the natural world, but also to devise and publish a diagram that united the notion of the “ladder” with the model of an intertwining genealogical tree. In his two-volume Philosophie zoologique of 1809, Lamarck encouraged his readers to visualise the emergence of a wide range of animal groups taking place as suggested in a branching diagram (Fig. 5). As this was intended to be read not as a model of development occurring from the bottom up, but from the top down, starting with worms (“vers”) at the upper left, it effectively maintained the model of the arbor porphyriana. And, as it privileged no particular starting point of development towards its upper edge, but proceeded in a zig-zag fashion down the page, it can also be seen as incorporating the double-trunked tree of Pallas’s model of nature, while at the marque dans l’arbre généalogique: l’influence du climat paroît y avoir produit les principaux changements, puisqu’on les observe également dans les deux autres races d’Amerique” (Duchesne 1766, Vol. I, p. 225). Cf. Zimmermann 1953, p. 257. 14  Berns 2000; Demandt 2002, pp. 19, 174–76.

13

14

II. From tree to coral (1837)

same time drawing on the “scale” proposed by Bonnet (Fig. 4). According to Lamarck, the device of merging the “scale”, or “ladder”, model with that of the “tree” allowed one to recognise “[that] the ladder corresponding to animal life has its beginnings in at least two separate branches, and that a number of shorter branches appear to have simply brought it to a halt”.15 The contrast bet­ ween change and stasis, indeed abrupt termination, is already to be found within Lamarck’s aforementioned positioning of worms at the upper left in his scheme and his grouping to the right of infusoria, polyps and radiata. While these last remain stable, and thus result in no subsequent divergence, the worms are seen to emit a diagonal dotted line, which in due course bifurcates, leading to the insects, arachnids and crustaceans, group­ ed on the right, and to the annelids, cirripedes and molluscs, positioned somewhat further down on the left. From this last group a further diagonal dotted line leads on to the fish and reptiles. These in turn are linked by further dotted lines to birds and then to monotremes (on the left) and to amphibians (on the right). From the last, in turn, diagonal dotted lines lead to cetaceans (to the right) and to both ungulate and unguiculate mammals (respectively, at the right and left termini of a final bifurcation). Lamarck’s hybrid of the ladder and the branching trunk of a tree befits the compromise at the heart of his own model of nature. In this model each group within the animal kingdom (most of the categories named there correspond to what would now be termed a phylum), and thus by implication each species, evolves in

4 _ Charles Bonnet, Idea of a Scale of Natural Beings. Illustration in his twovolume publication of 1764–65, Contemplation de la Nature

15  “[que] l’échelle animale commence au moins par deux branches particulières, et que, dans le cours de son étendue, quelques rameau[x] paraissent la terminer en certains endroits” (Larmarck 1809, Vol. II, p. 462).

1. The tree model

5 _ Jean Baptiste Lamarck, Diagram of the emergence of distinct animal groups. Illustration in his two-volume publication of 1809, Philosophie zoologique

accordance with its own inherent will to perfection; but, in doing so, it never transgresses its appointed limits.16 In order to convey, simultaneously, the notion of stasis and that of change, Lamarck’s diagram functions here through the interplay of text and image.17 While the arrangement of the named phyla – essentially balanced to left and right of an implicit central “axis”, albeit avoiding strict horizontal alignment – supports the static model of the ladder, the diagonal dotted lines originating with the worms at the upper left, emphasise the idea of development. Through notionally merging the rungs of a ladder with the new growth sprouting from a tree, Lamarck succeeded in capturing and communicating his principle of “willed” transmutation in a puzzle picture that would lend support – depending almost entirely on the stance of the viewer – both to the idea of a natural world unchanging since the Creation and to that of a world of living forms engaged in perpetual evolutionary growth. 16  Junker and Hossfeld 2001, pp. 49–57. The linear interconnection that Lamarck assumed to exist between the organs of digestion, respiration, movement, reproduction, and sensation (the nerves), and circulation reinforced the symbolism of stasis (Lamarck 1815, p. 367; cf. Métraux 2001, p. 16). 17  On the history of the diagram functioning, and considered, as an image: Bonhoff 1993; Bogen and Thürlemann 2003.

15

16

II. From tree to coral (1837)

2. Darwin’s first sketches In July 1837 Darwin set about tackling the compromise that, in his own view, fatally undermined the Lamarckian model. The first, pen-and-ink sketches that he drew in the aforementioned Notebook B (Fig. 6), apparently shortly after invoking there the model of the branching tree, nonetheless reveal that he had found inspiration in Lamarck’s use of dotted lines. Darwin’s rapidly drawn forms were to become the starting point for several decades of reflection. The very first of these (Figs. 6, 7) comprises, in its lower half, just such a linear sequence of dots, which in due course fans out to form three dotted lines, each of

6 _ Charles Darwin, First and second sketches of evolution (species transmutation) through a process of “natural selection”. Pen and ink on paper, 1837. Cambridge University Library, DAR 121: Notebook B, fol.  26r. Reproduced with the permission of the Syndics of Cambridge University Library and of Willaim Huxley Darwin 7 _ Charles Darwin, First sketch of evolution. Detail of Fig. 6 8 _ Charles Darwin, Second sketch of evolution. Detail of Fig. 6

which soon becomes a continuous line. As revealed by Darwin’s observation, on an earlier page of Notebook B, the three continuous strokes are intended to indicate species diverging in response to distinct natural environments, dominated respectively by water, earth and air.18 Darwin’s crucial innovation here lies in his distinction between dotted and continuous lines. While the latter are here intended to represent species that are still living in each of the three natural realms (of water, earth and air), the dotted lines, both of the single “trunk” and of the radiating “branches”, represent those that are already extinct (which had lived, that is to say, in an era that could now be recon18  Cambridge University Library, DAR 121: Notebook B, fol.  23r. Darwin here writes of the “triple branching in the tree of life owing to three elements air, land & water”. Cf. note 2.28.

2. Darwin’s first sketches

structed only through what was to be learnt from the fossil record). On the page of Notebook B that follows his sketches Darwin observed: “We may fancy [,] according to shortness of life of species [,] that [,] in perfection, the bottom of branches deaden[s]”.19 Through this express differentiation between continuous and dotted lines, Darwin armed his epoch-making protest against Lamarck with the fire-power of the two simplest elements in geometry. For the first time, the transmutation of species – evolution, as this term is now commonly understood – had assumed a visually appreciable form. To the lower right of Darwin’s initial sketch is to be found a smaller drawing in which a short dotted line bifurcates into two continuous lines. The observation – in

fact, a question (albeit the question mark is missing) – written beneath the first sketch (Fig. 6, lower half of sheet) suggests that the smaller figure relates to the descent of all types of fish from a single, simple organism.20 Here, too, the initial use of a dotted line is intended to represent living forms that are now extinct. In Darwin’s second sketch, at the bottom of the same page of Notebook B (Figs. 6, 8) he seeks to convey, in a diagrammatic guise, the greater complexity of the descent of birds in relation to that of fish.21 This sketch exploits in full the pos19 

Cambridge University Library, DAR121: Notebook B, fol.  27r. Cambridge University library, DAR121: Notebook B, fol.  26r: “Is it thus fish can be traced right down to simple organization – birds not [?]”. 21  Gruber 1974, p. 136. 20 

17

18

II. From tree to coral (1837)

sibilities afforded by the use of both dotted and continuous lines. From the starting point a dotted line, signifying extinct living forms, now traceable only through fossils, curves up to the left, while on the right a somewhat shakily drawn continuous stroke signals those still living. Darwin’s advance upon Lamarck’s combination of “ladder” and “tree” models is clear in this sketch, too. When Darwin employs his dotted lines exclusively for those creatures still preserved only in fossil form, a far deeper sense of time and the passage of time enters Natural History. And one may, with some justification, claim that the zoologist is thereby transformed into an explorer of time as manifest in the geological record. Darwin’s sketches harbour a scientific and cultural-historical significance that it would be hard to over-state: they are unprecedented in treating the “tree of life” and the “tree of nature” not as vener­able inherited mental models, but as conventions (by no means unproblematic in themselves) for addressing the phenomenon of a process taking place in real time.22 As a rupture with the account of Creation found in the Old Testament, Darwin’s stance here truly resembles that of Copernicus when he dared to argue that the Earth might in fact no longer be able (through man, its “noblest” inhabitant) to lay claim to an astronomically “central” position. Admittedly, both of these 1837 notebook sketches might well be dismissed as negligible in purely aesthetic terms; and Darwin, throughout his life, greatly regretted that he did not possess the talent for draughtsmanship that distinguished the work of friends such as Joseph Hooker.23 By the same token, it appears all the more impressive that he nonetheless strove to devise a visual formulation for what was to prove a conceptual turning point. The result bears witness to the fact that Darwin’s perhaps most risky idea arose out of a tentative interplay between notes and sketches, and that these last, in spite of their ungainly form, were in themselves profoundly compelling.

3. The model based on coral On account of their bi- and trifurcations, both of Darwin’s sketches are suggestive of an abstract rendering of a “tree of life” of the sort that he had already proposed as a model for what was to become his mature concept of evolution. At the same time, however, the analogy seems not quite right in as far as the entity shown in the left half of the lower drawing (Fig. 8), with its dense cluster of shoots, more closely resembles a bush. And the resemblance is in fact more telling than it might at first seem. For what is improbably spectacular about this drawing derives not only from its apparent status as one of Darwin’s earliest visual representations of his future 22  On Darwin’s predecessors in infusing Natural History with the element of time: Lepenies 1978, pp. 41–43, and Bredekamp 2000, pp. 73–76. 23  Darwin: Autobiography (ed. 1958), pp. 77–78.

3. The model based on coral

9 _ Petrarch Master, The Tree of Man. Wood-cut illustration to 1532 German edition of Petrarch’s collection of dialogues (1350), De remediis utriusque fortunae

theory, but also from its evidence of his effective re-working of the tree model as an alternative form of visual symbol. It is clear that Darwin, at the point where he sought to render, in a sketch, the general pattern of evolution in nature, as he was beginning to understand this, was well aware of the weaknesses of the tree as a metaphor – weaknesses that had, indeed, repeatedly incurred dissatisfaction since the Mediaeval period. This had above all been on account of the manifestly uni-directional growth of such metaphorical entities, which were essentially derived from those genealogical trees that served to characterise and rank an entire society, and in whose uppermost branches only those of the noblest birth were to be found. A satire on the tree model in this capacity was included, by the so-called Petarch Master, among his wood-cuts illustrating a 1532 German edition of Petrarch’s own text of 1350, De remediis utriusque fortunae, a series of 254 dialogues on consolation for the vagaries of fortune. Here (Fig. 9), the

19

20

II. From tree to coral (1837)

peasants glumly stationed among the roots at the base of the tree are again encountered, luxuriating in leisure, in its highest branches; and that on the left, seen playing a set of bagpipes, may even be found to have his feet planted on the tip of the mitre and on the sumptuously robed torso of the figure of a pope who sits on a slightly lower branch.24 In the realm of the Natural Sciences, meanwhile, the Italian archaeologist and botanist Vitaliano Donati had proposed, in 1750, as an alternative to models based on a ladder or a tree, that it might be possible to develop a system for ordering knowledge of the natural world through the use of a model based on a network of knotted threads, and that this might well prove far more “elastic”, and hence usefully accommodating, than would any attempt at a linear presentation of the complexities likely to be encountered: “In each one of the orders or classes derived in this way, nature brings forth her [own] sequences, passing imperceptibly from one circle to the next, as if along an interlinked chain. Be that as it may, the circles of a chain are nonetheless so tightly bound with each other that one would do well to compare natural progression less with a chain than with what might be called a net, for the latter is woven out of a number of threads, which in combination are found to evince complementary forms of communication, correlation and union”.25 One might argue that it is very much in the tradition of this negative critique of the tree model that Darwin, on the notebook page that directly precedes that bearing his two small sketches, made the following assertion: “The tree of life should perhaps be called the coral of life”.26 And his further, elucidating comments reveal that, at this phase of his thinking on the matter, his use of the term “perhaps” was merely rhetorical. It was in particular with regard to those parts of the coral formation that were no longer living that Darwin would have been able to reject Lamarck’s notion that “willed” transmutation occurred in a continuous process, in favour of a drama that truly was a matter of survival or death. His observation on “the coral of life” in fact goes on: “[…] base of branches dead; so that passages cannot be seen. – this again offers contradiction to constant succession of germs in pro­ gress”.27 Darwin would have favoured coral as a model because, with its petrified 24 

Berns 2000, pp. 156–57. ”In ciascheduno di tali ordini, o Classi la natura forma la sua serie, ed ha li suoi insensibili passaggi da anello in anello delle sue catene. Oltre di che gli annelli d’una catena talmente sono uniti con quelli d’unaltra, che ad una rete piuttosto, che ad una catena le naturali progressioni si dovrebbero rasomigliare, essendo, per dir così, tessuta di vari fili, che tra loro hanno scambievole communicazione, correlazione, ed unione” (Donati 1750, p. XXI). Cf. Métraux 2001, p. 10. 26  Cambridge University Library, DAR 121: Notebook B, fol.  25r. 27  Cambridge University Library, DAR 121: Notebook B, fols. 25r–26r. On this point, cf. Hemleben 1979, p. 94; Pörksen 1998, pp. 327–28; and Treusch-Dieter 1994, who, taking into account material other than that considered in the present volume, see in these comments of Darwin’s a starting point for an understanding of his work that encompasses even his last major work, on the expression of the emotions in man and animals (Darwin 1872). 25 

3. The model based on coral

10 _ Corallium rubrum. Berlin, Museum für Naturkunde, Cni 352

stem (which could be seen to stand for the fossil form of an extinct species) and its radiating limbs, it could offer a counter-image to Lamarck’s notion of continuous transformation. The words inscribed along the top of the notebook page bearing Darwin’s two sketches (Fig. 6) – “no only makes it excessively complicated” – would seem to function as a means of pre-empting potential objections to his own aversion to Lamarck. In coral Darwin found a model of evolution as he would in due course come to understand it, a model that made it possible to visualise the chronological process more powerfully than did the tree model because it made clear, at a glance, the separation of species still living from those that were already dead. The dotted lines used in Darwin’s sketches have a loose, but more than merely incidental, equivalent in the stubs ranged along the petrified limbs of many sorts of coral formation – or, indeed, in the distinct “bosses” of Darwin’s Patagonian specimen (Fig. 1). In making his first sketch (Figs. 6, 7), Darwin might well be seen to have had in mind a type of coral found in the Mediterranean: Corallium rubrum (Fig. 10). With its emphatically bifurcating structure, it is close to the form evoked on the notebook page. While such a form might, of course, have been suggested by that of a deciduous tree deprived of all its leaves in winter, the aforementioned coral may well have held a particular fascination for Darwin because it appeared to embody, in an exemplary fashion, one of the central aspects of his critique of Lamarck. While the latter’s concept of “willed” transmutation assumed that any change occurring, on whatever scale, would always be restricted to that natural realm (water or earth

21

22

II. From tree to coral (1837)

or air) to which the living form in question belonged, change as envisaged by Darwin assumed that living forms (more specifically, particular species) might, on occasion, attain the capacity to “breach” the limits of their original habitat. As Darwin asked: “Would there not be a triple branching in the tree of life owing to those elements air, land & water; & the endeavour of each typical class to extend his domain into the other domains & subdivision three more, double arrangement. –”.28 Just as Darwin, even in his earliest notebook allusion to a branching tree, had implicitly taken issue with Lamarck, so too does his reference to a notional doubling of the three realms of nature reiterate his view that each living form may seek to transgress the limits of its own habitat, be this in the water, on land or in the air. In yet another annotation he had observed: “[…] if each main stem of the tree is

11 _ Melitodes flabellifera var. cylindrica. Berlin, Museum für Naturkunde, Cni 5437

adapted for these three elements, there will be certainly points of affinity in each branch”.29 And, even in Darwin’s subsequent summation of his thoughts on the process of creation in nature, he was to emphasise: “[…] every part would probably not complete [i.e. attain completion], if birds were fitted solely for air & fishes for water”.30 It was, however, precisely this sort of potential for interconnection that was not easy to visualise with an arboreal form; for the clearly distinct branches (which in Darwin’s sketches symbolise the three realms of nature) do not, in the case of a tree, go on to intertwine. With a coral formation, by contrast, even when those parts that might be viewed as the “branches” and “twigs” have grown far beyond their respective budding points, they are nonetheless entirely capable of forming new, cohesive units (Fig. 10). As a result, coral assumed, for Darwin, an anti-Lamarck28 

Cambridge University Library, DAR 121: Notebook B, fol.  23r. Cf. note 2.18. Cambridge University Library, DAR 121: Notebook B, fol.  24r. 30  Cambridge University Library, DAR 121: Notebook B, fol.  45r. 29 

3. The model based on coral

12 _ Leptogorgia petechizans. Berlin, Museum für Naturkunde, Cni 4836

ian symbolism. Combined with its capacity to exhibit the extinct species as a petrified stem, this could not but entrance him. Darwin would appear to have had such a form in mind when devising the figure on the left of his second sketch (Figs. 6, 8) in so far as its projecting shoots are rendered in a manner more suggestive of a bush than of a tree. A bush-like appearance of this sort is a characteristic shared, for example, by those similarly extending and then again interlocking limbs of Melitodes flabellifera var. cylindrica (Fig. 11) or the fan-like fronds of Leptogorgia petechizans (Fig. 12). Among the “Corallinae” amassed by Darwin when he was on the H.M.S. Beagle expedition are to be found several organisms that have a tendency to evince the bush-like appearance captured in Darwin’s second sketch. Among these finds, three examples of Corallina officinalis (Fig. 13), a further coralline alga that Darwin at that time took to be a coral,31 may be seen, on account of their dense dishevel31 

pact”.

Darwin: “Coralline Algae Notes” (ed. 1987), p. 192: “Corall exceedingly hard, stony com-

23

24

II. From tree to coral (1837)

13 _ Corallina ­offi­cinalis. London, Natural History Museum, Darwin 1143

ment, as an extreme form. No sooner has each shoot advanced a little distance from the stem than it begins to intertwine with others. Examined in greater detail, moreover, each ostensible “stem” is found, as Darwin noted, to be “composed of numerous small oblong pieces, with globular heads, these often grow into each other & are always close together”.32 (And one is here reminded of the tightly aligned “bosses” of Darwin’s Patagonian specimen, Fig. 1). It is this sprawling structure that is to be found in a yet more emphatic form in the third pen-and-ink sketch that Darwin made, in 1837, in Notebook B (Fig. 14), from which anything resembling a tree has vanished. In order to affirm this definitive shift in both script and image, Darwin placed at the upper left of the page what might now almost be understood as a sort of epigraph: the words “I think”. When the drawn form was added, the inscription could well have been seen as characterising this as itself a membrane for thought.33 In retrospect this has come to be recognised as one of the most happily inspired moments in the History of Science. Darwin’s image is neither a derivation from nor an illustration of something observed in the natural world. It is, rather, an active bearer of the process of thinking. It is as if the words “I think” are inscribed by the thinker, but simultaneously spoken by his sketch. What is striking about the form that Darwin here drew is that its proliferation does not appear to have occurred in response to the largely upward “striving” one would associate with a tree, but rather to reflect an impulse to send out new growth in every possible direction. A line, identified at its base with an encircled “1”, 32  33 

Darwin: “Coralline Algae Notes” (ed. 1987), p. 192. On this point: Voss 2003 Darwins Diagramme, pp. 1–2.

3. The model based on coral

14 _ Charles Darwin. Third sketch of evolution (species transmutation) through a process of “natural selection”. Pen and ink on paper, 1837. Cambridge University Library, DAR 121: Notebook B, fol.  36r. Reproduced with the permission of the Syndics of Cambridge University Library and of William Huxley ­Darwin

ascends a short way up the page, then divides into three: a shoot to the left comes to a rapid end, while that maintaining the initial vertical thrust and that extending to the right appear to thrive, the former itself soon bifurcating to both left and right, and these shoots in turn doing likewise, the latter giving rise to a series of further divisions. According to the summary explanation that Darwin has added in the lower half of the page, each of the letters A, B, C and D represents a variety of the same species, the distance between the letters serving as a measure of their divergence: “Thus between A & B. immens[e] gap of relation. C & B. the finest gradation, B & D rather greater distinction” At the upper right of the page Darwin has added the following deduction: “Case must be that one generation then should be as many living as now”. Perhaps significantly, these words may be read either as a continuation of the initial use of “I think” at the upper left, or as a free-standing observation, or even as both at the same time. The text at the upper right treats, in fact, of the theory of the monadic balance of the emergence and extinction of species, as a result of which the number of those that momentarily prevail in the struggle for survival will always be matched

25

26

II. From tree to coral (1837)

by the number of those defeated in this struggle, meaning the overall number of living species remains constant.34 The subsequently encased smaller segment adds further enlightenment: “To do this & to have many species in same genus (as is) requires extinction”. A renewed examination of the sketched form itself reveals the extent to which Darwin has also articulated this “balance sheet” of survival and extinction in graphic terms. The central stem marked “1” may be found to have given rise, both directly and at one or more removes, to 25 varieties. However, only half of these can survive; and, as 13 here appear to be stable, 12 would, over time, have to become extinct, as Darwin observes on the very next notebook page.35 In order to render this genealogical mechanism in visual form, Darwin has distinguished surviving species through the addition of a short bar placed at right-angles to the extremity of each of the respective lines. This is the case with all three shoots grouped under “A” and under “C”, as also with two of the three grouped under “D” and all four grouped under “B”: a total of 13 living species. These are “balanced” by 12 extinct species. The latter are here shown as shoots that merely terminate without the addition of a cross-bar: the four that extend from the rightward curving stalk above “A”, and a sub-total of five to the left and three to the right of the initial vertical shaft. The form of Darwin’s sketch was to some extent prompted by his interest in the diagrams made by the Scots physician Martin Barry, who in early 1837 had introduced the British scholarly community to the theory of archetypes put forward by the Baltic German embryologist Karl Ernst von Baer.36 Darwin’s own drawn thought-experiment recalls Barry’s proposal that both men reject a more elevated understanding of evolution, as an ever closer approach towards “perfection”, in favour of the notion of a mechanism of differentiation. And, while Barry’s Tree of Animal Development (Fig. 15) seems, at first glance, to have little in common with Darwin’s sketch, it is in this context significant that, in both, the overall appearance in no way evokes the image of a tree in nature. By contrast with Barry’s crystalline and “prickly” branching forms, however, Darwin’s own model is less assertive, more irregular, more organic; and, on account of these characteristics, it may be said to take its bearings far more than do any of Darwin’s other early sketches, from the structure of coral.37

34 

Gruber 1974, pp. 143–44. Cambridge University Library, DAR 121: Notebook B, fol.  37r: “[…] for if each species […] is capable of making 13 recent forms, twelve of the contemporarys must have left no offspring at all, so as to keep number of species constant. – ”. 36  Barry 1836–1837, p. 346. This was first observed in Voss 2003 Darwins Diagramme, pp. 23–25, for which see also what follows. On the influence of Baer on the arboreal model: Ospovat 1981, pp. 158–59 and passim. On Baer: Breidbach 1999. 37  Voss 2003 Darwins Diagramme, p. 26. 35 

3. The model based on coral

15 _ Martin Barry, Tree of Animal Development. Illustration (detail) in his article published in the April 1837 issue of the New Edinburgh Philosophical Journal

In later years Darwin never ceased to employ this form of graphic model. The verso of an undated advertisement for the services offered by a printer, Edward Strong, bears what appear to be three distinct layers of script and diverse marks (Fig. 16). In the earliest of these there are statements, single words and sketches, all in pencil. The layer above this is dominated by four rounded, interlocking shapes, all bearing text in what is now dark brown ink. Pencil is again used in a third, less crowded layer, both for script and for further sketches. The passage along the top of the sheet bears the pencilled allusion to a “shortfaced & long-faced Tumbler” (a “tumbler” being a variety of domestic pigeon) and, below this, a trifurcated form labelled “Descent” at its base. Below this sketch, in turn, and within a funnel-shaped bubble that narrows to the right, is to be found a spiky confection with bifurcating lines, the meaning of which is revealed in an inscription placed outside the bubble towards the right of the sheet: “This is explanation of longer / greater similarity of Embryo of Pig & Cow”. By contrast with these two sketches in the upper left section of the sheet, with their characteristic bifurcating structure, one finds, just a little below the centre of the page, an image that seems to emerge from a different intellectual context. Dar-

27

28

II. From tree to coral (1837)

16 _ Charles Darwin, Sketches relating to descent and embryonic de­velopment, on verso of an advertisement of services offered by the printer Edward Strong. Pencil, pen and ink on paper, probably 1840s. Cambridge University Library, DAR 205.6: portfolio of loose sheets, fol.  51. Reproduced with the permission of the Syndics of Cambridge University Library and of William Huxley Darwin

win has drawn its lines in short, nervous segments, as if he were seeking to coax into cohesion a wafting entity that is not altogether willing to stand erect, preferring to creep, in a tangle, along the ground. This contrary character is all the more striking in that it is in fact concerned with animals all belonging to the same family, their names displayed, on the left, as if upon a barbed frame: “cat”, “dog” (twice) and, finally, “hyena”. The most complex sketch, drawn in pencil across the upper half of the sheet, is partially obscured by Darwin’s subsequent writing in ink and the lines drawn around its separate segments (Fig. 17). As indicated by the word “Mammals” that appears at its base, it concerns their evolution, which is indicated in a sequence of lines extending both vertically and diagonally to both left and right. Already hard

3. The model based on coral

17 _ Detail of Fig. 16.

to decipher through the overlay of ink lettering, the drawn line furthest to the right pursues a markedly indirect path. At its highest point it appears to take a sharp turn to the right and then to engage in an essentially “downward” sequence of bi- and trifurcating manoeuvres – a pattern already to be found in the passage from “1” to “A” in the third coralline sketch of 1837 (Fig. 14). It is no coincidence that this, apparently the first sketch to be made on this sheet, is the one most closely resembling the structure of coral. It is, moreover, also the one that best exemplifies Darwin’s grasp of the interaction of regularity and chance that he had perceived to be at work in the sheer profusion and differentiation of the forms found in nature.38 Not only could coral convey the compelling image of evolution as a sort of universal “battle painting”, in which the living victors prowled about among the petrified dead. On account of the pattern of its growth it could also represent the anarchic aspect of evolution – and thereby reveal as erroneous a mimetic understanding of this, such as was invariably encouraged by the model of a tree. Finally, with a vigorous pencil line, looping in several stages from left to right, Darwin has circled each of the three most obviously coralline passages within his sketch, thereby also enhancing the significance of its position at the centre of the sheet. The element of trial-and-error in this image-and-thought experiment, as also the eventual palimpsestic character of the document as it has been preserved, make 38  Darwin has already referred, employing a surely deliberate oxymoron, to the “law of chance”. See: Cambridge University Library, DAR 121: Notebook B, fol.  55e. This, in his view, stood in opposition to what he was beginning to understand of the “laws” of evolution.

29

30

II. From tree to coral (1837)

of this an especially striking testament to the character of Darwin’s procedure. It would be hard to find as succinct an encapsulation of his own understanding of nature in its entirety as is to be discovered in this at first unpromising series of sketches. It is through their efficacy and versatility as a means of posing questions and testing hypotheses that we encounter Darwin on the track of a convulsively variable natural world.

III. Strickland’s alternatives (1840)

1. A critique of metaphor Darwin’s search for models of a natural world characterised by variability was taking place in parallel with the efforts of numerous comrades-in-arms. It is, indeed, probable that in few other phases of the History of Science has the devising of models been so fervently debated as it was in England between 1835 and 1860, the very years during which Darwin was thinking his way towards his own theory of evolution. Two questions – whether each of the species could be treated in relation to a corresponding natural system or might be merely arbitrarily defined, and whether the process that Darwin was to identify as “natural selection” had occurred as continuous change or in a series of sudden leaps forward – were central to the competing theories. Prominent among those of Darwin’s contemporaries who took a similar interest in metaphors for the natural world was the now largely forgotten geologist and zoologist Hugh Edwin Strickland, who in 1853 was to meet an early death, at the age of 42, in a manner no less bizarre than it was tragic: he was crushed under an approaching, but unheard, locomotive while hunting for fossils in a railway cutting.1 Strickland’s historical significance here lies in his prolonged effort to establish a systematic method of naming species. In the dispute between those who still maintained that the diverse species had been created in accordance with a Divine Plan, and had thereby each also acquired its name, and those who had become convinced that the existing designations of both genera and species were abstract formulations with no direct correlate in the natural world, Strickland insisted that the human act of naming or definition must be transparent – the reasoning behind it remaining clear.2 Darwin’s own scepticism regarding the fixing and naming of species had been intensified through his conversations with Strickland, with whom he remained in contact throughout the 1840s, and whose organisational assistance 1  2 

Browne 1995, p. 451; fundamental on this matter: Di Gregorio 1987. McOuat 1996, pp. 511–15.

32

III. Strickland’s alternatives (1840)

he sought, and received, in his attempts to outmanoeuvre opponents.3 When considering who might prove to be the best editor for his work in the event of his death (prompted to do so by having, in 1844, committed to paper a first full summary of his theory), Darwin included Strickland among those he thought most suited to this role.4 Darwin had been particularly impressed by Strickland’s article of 1840 in favour of the visual representation of the interrelations, within a given large animal group, between orders, families, genera and species; and this is, indeed, still recognised as a milestone in the use of metaphors in Biology. Convinced that nature, in all its disorderly profusion, could not be constrained within a rigid framework and a systematic scheme, Strickland was enraged at every attempt to make the natural order more easily comprehensible through metaphors entailing chains, circles, networks, or symmetries based on number theory.5 In disallowing even the notion of a “great chain” of living creatures – the idea that each species possessed only two neighbours, one lying to either side of it – Strickland was eschewing a concept that had been widely accepted since the early years of the nineteenth century.6 While such a “chain” could, of course, be nominally subdivided into endlessly differentiated segments and intermediary links, Strickland nonetheless rejected it on account of its inextricable connection with the notion of entities arranged in a single straight line. Strickland was no less dismissive of the circle-based models applied to the natural world by the entomologist, and acclaimed expert on beetles, William Sharp Macleay, and popularised by, among others, the ornithologist William Swainson.7 The Horae Entomologicae, published by Macleay in 1819 and 1821, had presented diverse aspects of the animal kingdom as arranged in five contiguous circles (Fig. 18).8 In December 1843, emboldened by Strickland’s negativity towards such models of nature, Darwin ironically dismissed Macleay’s system of “vicious circles”.9 3  See, for example, the intensive correspondence of January–February 1849, largely on issues (of great interest to Darwin at this time) of taxonomic nomenclature: Darwin: Correspondence, Vol. IV (1988), nos. 1215, 1216, 1221, 1223, 1224, 1225, 1226, 1227, pp, 187–218. On organisational co-operation with Darwin: McOuat 1996, p. 507. 4  See Darwin’s memo of 5 July 1844, to Emma Darwin, in Darwin: Correspondence, Vol. III (1987), no. 761, pp. 43–44, here 44. The 1844 manuscript is now in Cambridge University Library, DAR7: “sketch of species theory, 1844”. 5  Strickland 1840, pp. 186–88. 6  Lovejoy 1936, passim. 7  Swainson 1836–1837, vol. II, 1837, p. 200; cf. Di Gregorio 1983, pp. 234–36; Ritvo 2003, pp. 60–62 and Voss 2003 Darwins Diagramme, pp. 15–16, also of relevance to the following discussion. 8  Swainston 1985, pp. 14–15; Di Gregorio 1987, p. 37; Gardiner 2001, pp. 7–9. 9  Letter of 3 or 17 December 1843 from Darwin to George Robert Waterhouse, in Darwin: Correspondence, Vol. II (1986), no. 718, pp. 415–16, here 416: “I believe infinite harm has been

1. A critique of metaphor

18 _ William Sharp Macleay, Diagram of the System of the Animalia, in Vol. I, part 2 (1821) of his Horae Entomologicae; or, Essays on the Annulose Animals

For Strickland himself, even the non-hierarchical arrangement of knotted threads, as implicit in his own concept of the natural world as a form of network,10 was in fact unsuited to encompass the “variety” of nature. “In short”, he wrote in 1840, “variety is a great and a most beautiful law of Nature; it is that which distinguishes her productions from those of art, and it is that which man often exerts his highest efforts in vain to imitate.”11 This “variety” in nature, here invoked as a principle in opposition to all the “laws” that govern art, derives from the tradition established by William Hogarth’s volume of 1753, The Analysis of Beauty, which had upheld

done by these circles, which catch the eye as of equal size, & inevitably lead the mind to suppose they are of equal value – it is by this artifice, as I believe, [that] the possibility of making the Quinarian system [= based on groups of five] appear probible [sic.] has chiefly rested: Moreover it shd be stated by everyone, I think, who indulges in these vicious circles, that confessedly there is no standard to judge of [= standard by which to judge] the value of groups. –”. Cf. Gardiner 2001, pp. 12–13. Cf. Strickland 1840, pp. 186–88. 10  See above, p. 28, for Donati’s earlier proposition of a network model. 11  Strickland 1840, p. 187.

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III. Strickland’s alternatives (1840)

the notion of the infinite variety of forms in nature in an anti-classicising spirit.12 In view of the irregularity to be discovered throughout the natural realm, rendering altogether hopeless every attempt at orderly representation, be it in a model or in a museum, Strickland concluded that all such systems were themselves a product not of nature but of art(ifice).13

2. From tree to map Such negativity on Strickland’s part did not, however, mean that he had rejected every form of visual metaphor. He too, indeed, had taken his starting point in the tree model. But, much in the spirit in which Darwin was to distance himself from the notion of the “tree of life”, Strickland was also moved to point out the weaknesses of such a metaphor. In his quest for a way to encompass the anarchic shaping power of nature, Strickland did embrace tree models that were sufficiently suggestive of irregular patterns of growth, even though eventually replacing the metaphorical notion of the tree with that of the ramifications of a primeval forest of unkempt bushes: “The natural system may, perhaps, be most truly compared to an irregularly branching tree, or rather to an assemblage of detached trees and shrubs of various sizes and modes of growth”.14 Even though Strickland is now only rarely identified as the first to invoke this metaphor, the image of the disorderly, bush-like tree has never entirely gone out of intellectual fashion.15 Finally, however, Strickland progressed to visualising the interrelationship of the species as a form of map, on which the representation of vegetal branching was no longer intended as a model relating to external forms, but as a pointer to a recommended methodology. To this way of thinking, the drawing of a tree on a page is no longer of significance on account of its formal properties, but on account of the notional positioning, on a two-dimensional planar surface, of a complex, threedimensional entity – of the application, that is to say, of the methods of cartography: “And as we show the form of a tree by sketching it on paper, or by drawing its individual branches and leaves, so may the natural system be drawn on a map”.16 The irregular distribution of species that Strickland found in nature was here opposed to the model of the tree, with its hierarchical arrangement of trunk, branch and twig: “The affinities of animals may possibly be neither rectilinear nor circular, but irregular like the distribution of water and land, and like that, capable of being 12  Hogarth 1753, pp.16–17 (chapter two: “Of Variety”). Hogarth and his Times, pp. 168–69; on this point in general: Gerlach 1989. 13  “[…] the design of creation was to form not a cabinet of curiosities, but a living world” (Strickland 1840, p. 187). 14  Strickland 1840, p. 190. 15  Illies 1983, p. 108. 16  Strickland 1840, p. 190.

3. The cartographic model

laid out on a map – not however by any a priori theory, but by long observation like [= as in] a geographic survey.”17 In favouring a cartographic approach, Strickland effectively shifted the problem of understanding the system of nature to that of its representation. In his view, one should look to the biologists not for a system truly suited to explaining what was to be found in the natural world but, rather, for a systematisation of the ways of representing nature’s own irregularity. Strickland’s endeavours to achieve an appropriate way to visualise nature were above all of value for this methodologically incisive conclusion. And it is in this that his thinking and that of Darwin coincided.

3. The cartographic model Strickland was far too critical a spirit not to admit that there were also serious weaknesses to be found in thus transferring the model based on the tree or the bush to the sphere of cartography. He was, nonetheless, prepared to remain optimistic regarding his invention: “[…] let us hope that the affinities of the natural system will not be of a higher order than can be expressed by a solid figure [,] in which case they may be shown with tolerable accuracy on a plain [= plane] surface, just as the surface of the earth, though an irregular spheroid, can be protracted on a map”.18 Strickland proposed a cartographic form of representation to convey the “natural affinities” among the “class of birds”. This showed the diverse groups and subgroups as if they were distinct land masses drawn on a map, in the manner of separate islands each surrounded by seawater. Strickland treated the spaces in a dynamic fashion, in each case determined by the interrelationships prevailing there. These are also suggested through the use of lines – although their strut-like character betrays its origin in the branching of irregular trees and of bushes.19 On 21st September 1840, in Glasgow, Strickland presented an expanded cartographic model of such “natural affinities” to an illustrious gathering of the British Association for the Advancement of Science (Fig. 19). This model demonstrated especially compellingly the interconnection of affinities (here registered through the presence of straight lines) with the irregular distribution of groups (here emerging like islands from an ocean).20 17  Handwritten note of 1838, cited after McOuat 1996, p. 501. On Strickland’s cartographic model and the history of its presentation, see McOuat 1996, pp. 501–04. 18  Strickland 1840, p. 190. 19  “In order to place these groups at their true distances, it is necessary to form a scale of degrees of affinity, to which the intervals between each generation shall correspond” (Strickland 1840, p. 191). The map is printed as plate VIII. Cf. Di Gregorio 1983, pp. 239–41; Di Gregorio 1987, pp. 40–41. 20  Jardine 1858, Fig. before p. cciv; cf. McOuat 1996, pp. 500–501, and Voss 2003 Darwins Diagramme pp. 13–14.

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III. Strickland’s alternatives (1840)

Strickland’s cartographic approach addressed the problem of how to represent aspects of the natural world in an appreciably systematic, and yet not over-simplifying, fashion. Through this approach he sought to transform the tree model, with its own irregularly mimetic relationship to the natural world, into a cartographic rendering of geographical contingencies. In proposing that the structural principles of cartography replace metaphors derived from the ladder, the tree, the network, or the circle, he was moved by the hope that nature might thereby be rendered in a manner not incompatible with “the wild luxuriance of her ramifications”.21 Darwin read Strickland’s 1840 article in the year of its publication; and the cartographic principle there introduced made so strong an impression upon him that he seems to have assumed that Strickland was its inventor. This does at least seem to be implicit in his comments of 1840 in his copy of the revised, 1836 edition of John Lindley’s Natural System of Botany (the first had appeared in 1830): “Does not Lindley [= Why doesn’t Lindley] use Diagrams to represent affinities, like the maps of Strickland?”22 Both editions, made up entirely of text, do indeed cry out for the addition of helpful and vivid diagrams, not least in the light of Lindley’s avowed concern with how a vast mass of factual data was best to be conveyed to readers.23 It is probable, moreover, that Strickland’s means of visualising the natural world held a particular attraction for Darwin because, in the three coralline sketches of 1837, Darwin’s goal had been not at all unlike that at which Strickland was aiming when he devised the model he had based on a bush and then the one he had based on a map. Darwin, in his sketches, had likewise sought to avoid the unidirectional character of the tree. And, were one to re-draft, in planar form, the respective positions of the terminal cross-bars used by Darwin to designate living species in his third 1837 sketch (Fig. 14), this would in fact produce a “map” such as Strickland was to envisage only a few years later. For all that they might be said to have had in common, Darwin and Strickland were, nonetheless, unalike in their respective approaches to the rendering of the dynamic adjunct of the passage of time. Strickland’s map of living creatures was a “snapshot” of species still in existence. In the third of the aforementioned 1837 sketches, however, Darwin sought, through tracking each living coralline limb in its distinct pattern of growth, to show the emergence of variation, and he found in the image and the notion of the petrified coral stem a compelling symbol for the realm of fossils. Herein lay the unmistakeable power of the coralline model that was to preoccupy him in the years to come.

21 

Strickland 1840, p. 192. Di Gregorio und Gill 1990, column 501, record Darwin’s comment in his copy of the second, 1836 edition of Lindley’s volume. 23  See Lindley 1830, pp. v–x (Preface). 22 

3. The cartographic model

19 _ Reduced version of Hugh E. Strickland’s chart of The Natural Affinities of the Class of Birds. Fold-out plate in William Jardine’s volume of 1858, Memoirs of Hugh Edwin Strickland

37

IV. From the circle dr awings to the diagr am (1851–1858)

1. The circle segments Three drawings made, in all probability, between 1851 and 1857 (all in pencil, two of them on the recto and verso of a single sheet) testify to Darwin’s renewed attempts to bring into play the element of time without thereby evoking the record of change of a teleological sort (Figs. 20, 23, 24).1 In their combination of image and text, these constitute Darwin’s further endeavour to find an appropriate way to visualise species transmutation as he approached his mature understanding of this. The off-white sheet (now yellow with age) used, as an upright rectangle, on only one side (Fig. 20), bears a complex arrangement of interconnected pencil lines, branching and re-branching as they extend from an emphatically marked point on the left, and occupying around 140 degrees of a circle, its circumference faintly traced just beyond their termini, with an even fainter inner circle visible shortly before these. Because the words inscribed directly to the left of the drawn structure – “Parent of Marsupalia and Placentalia” – are not placed more or less horizontally (as are the inscriptions to the right), they require anyone wishing to read them to turn the sheet through 90 degrees to the left. This then reinforces that reader’s readiness imaginatively to “complete” the here merely intimated full circle. And, as a circle admits of no hierarchy of direction, Darwin’s annular acknowledgement of the passage of time may be recognised as an endeavour to map chronological phases not merely so as to register growth in one direction (as would be the case in a model based on a ladder or a tree), but rather so as to trace its simultaneous occurrence in every direction. It is here as if the record of the ramifications of a bush-like tangle of shoots has been scratched on to the pattern of tree-rings revealed on the inner surface of a severed trunk. Darwin was here partially following the approach taken by a close, FrancoSwiss contemporary, from 1847 teaching at Harvard University, the geologist and zoologist Jean Louis Rodolphe Agassiz, in his 1848 volume, co-authored with the 1 

Voss (2003, Darwins Diagramme, p. 31) believes these drawings were all made in 1853/54.

40

IV. From the circle drawings to the diagram (1851–1858)

20 _ Charles Darwin, Drawing of evolution (species transmutation) through a process of “natural selection”. Pencil on paper, 1851–57. Cambridge University Library, DAR 205.5: portfolio of loose sheets, fol.  183r. Reproduced with the permission of the Syndics of Cambridge University Library and of William Huxley Darwin

American conchologist A. A. Gould: Outlines of Comparative Physiology. The frontispiece to this publication (Fig. 21), showing the Crust of the Earth as Related to Zoology, presents the globe through time as a circle comprising eleven concentric rings (each representing a geological era, and coloured according to the table below) around a central, inviolate point of origin, itself directly encased in a white shell bearing symbols of four “archetypes” (their names – molluscata, radiata, articulata, vertebrata – are found inscribed around the outermost ring). Beyond this white

1. The circle segments

21 _ Jean Louis Rodolphe Agassiz, Crust of the Earth as Related to Zoology. Frontispiece to enlarged, 1851 edition of his 1848 volume Outlines of Comparative Physiology

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IV. From the circle drawings to the diagram (1851–1858)

shell the various zoological categories are encountered, in the form of labelled white shafts, each burrowing outwards through the concentric chronological layers towards the two outermost circles, which for Agassiz constitute a bi-partite “Modern Age”. A few of these (for example, the “ammonites”, among the molluscata) are present at an early date but, as in this case, extinct before the advent of the “Lower Tertiary Formation”; others (for example, the “frogs”, the “whales” and “man”, all among the vertebrata) arrive only later but persist into the present era. On account of the wide readership reached by the Agassiz-Gould volume, its vivid introductory image did much to help establish the circular variant for models of evolution such as endures to this day. Its advantage lay in the fact that the increase in differentiation between the zoological categories was vividly rendered through the increase in the distance that intervenes between these as they move further and further from the centre, albeit this is immediately clear only in the case of the longest enduring entities (for example, the “brachiopods” and the “bryozoa”, among the molluscata).2 The frontispiece alerted Darwin himself to the possibilities afforded by showing the system of nature in the form of a circle. In his own work, however, he responded not by indicating the distribution of specific categories through the use of shafts but, rather, through rendering this distribution with his own scheme of radiating and bi- / trifurcating lines, thereby placing greater emphasis on differentiation and variation. In the aforementioned Darwin drawing (Fig. 20) the line that extends more or less vertically from the point of origin fans out expansively, though only after it has advanced quite a distance. By contrast, the neighbouring line, which ascends diagonally towards the upper right of the sheet, starts to divide almost at once, but does so far less vigorously or prolifically. This second shoot has been circled by Darwin, while a line also running to the upper right relates it to the words inscribed there: “no form intermediate”. The third line to emerge progresses more or less horizontally to the right, but is itself distinguished by soon emitting a far more delicate shoot to its own left. This in turn soon gives rise to an entire series of feathery sprigs, which Darwin labels (once within the main structure and once outside the circle) “Rodents”. Below this are two lines extending, almost in parallel, to the right: both are non-branching continuations of the early stage of the aforementioned third shoot, and one is labelled (at its extremity) “Marsupials”. In marked contrast to these examples of undiversified growth, is a final, highly prolific form running to the right and lower right. Darwin has here apparently wished to emphasize the possible close proximity of growths with the most complex and delicate ramifications to those that produce not a single sub-division.

2  A scheme similarly based on the segments of a circle was proposed in 1960 for the higher mammals (H. Schmidt 1960, p. 269); cf. also David Hillies’s Roundabout, in: Pennisi 2003, p. 1697.

1. The circle segments

If one now turns the sheet through 90 degrees to the left so that the aforementioned inscription “Parent of Marsupalia and Placentalia” becomes more easily legible, the logic informing Darwin’s drawing can be more fully appreciated. For it is only from this new perspective that it becomes apparent that the centre of the implicit full circle is located not at the point of origin of the branching structure here described, but at a point lying directly above the initial letter of the word “Placentalia”, which Darwin has, indeed, marked with a tiny dot (Fig. 22). A further

22 _ Detail of Fig. 20

dot stands to the left of the first, and a more faintly drawn broken line wanders to the left, only to terminate after a few segments. But a slightly firmer, dotted line runs diagonally to the upper left, traversing the word “Parent”, then merging with the ostensible base of the main structure. Attention to this most minute of details is in fact crucial to an understanding of this sheet as a whole. For it reveals that Darwin has here achieved a combination of the circular rendering of the system of nature, as devised by Agassiz, and his own earlier coralline sketches, which had designated extinct species through the use of dotted lines and those still living through the use of continuous lines. Darwin’s first coralline sketch (Figs. 6, 7) is, in most respects, identical to the small, further dotted-linear form in the later drawing, which runs vertically to the upper right of the word “Parent” before dividing into four (Fig. 22). Of these, all but the second line from the right are rendered in dots. Darwin finally accounts for this abundance of extinct coralline limbs in the words inscribed to the extreme left of the sheet (and requiring, for easier legibility, that it be turned back through 90 degrees): “if they had all given [= had] descendants then there w[ould] have been a great series”. Darwin’s preference for a combination of dotted and continuous lines reveals that, even in the present drawing, he was thinking not in terms of a tree model, but in terms of a model based on the form of coral. As this drawing also registers the passage of time through the presence of concentric circles (albeit rather faintly

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IV. From the circle drawings to the diagram (1851–1858)

drawn), it dispenses with a system of co-ordinates directed exclusively upwards or the indication of growth through parallel vertical lines. In Darwin’s drawing the ramifications tend not upwards, but outwards; and in this they correspond both to the “crust of the earth” model devised by Agassiz (Fig. 21) and to the model informing Strickland’s map (Fig. 19), both of which implicitly permit movement in any direction. Along the upper edge of the drawing Darwin has provided a further testament to how deeply he was preoccupied with the graphic elements that he had introduced decades earlier into his first coralline sketch. Characteristically, this is phrased as a question: “Let dots represent Genera???” It remains, however, unclear as to whether Darwin is here referring to the “dots” of the lines he had repeatedly used to designate extinct species, or to the small bubbles to be found at some of the termini of the drawn branches of the “Rodents” and the “Marsupials”. In this matter, however, the recto of the second, blue sheet (which also bears a drawing on its verso) is unambiguous on account of the definition supplied in its own inscription (Fig. 23): “Dot means new form – say in Birds.” By comparison with the drawing relating to the “Rodents” and the “Marsupials”, it is evident that a new system has been applied. Here, Darwin includes no dotted lines. Rather, he uses somewhat larger and emphatic dots as marks added to continuous lines, both subdividing these into several segments, and reinforcing those points at which divergence occurs. In addition, apparently following the model supplied by Agassiz (Fig. 21) – where a “Palaeozoic Age” constitutes the three innermost circles and around this are placed four rings representing a “Secondary Age” and two rings representing a “Tertiary Age” – Darwin labels his own concentric circles, from the centre outwards, “Palaeoz[oic]”, “Second[ary]” and “Tertiary”. The drawn lines here are fewer and, on the whole, straighter; and they are relatively evenly positioned in relation to the slightly less faint markings of the circles. In this case three very distinct outcomes are of particular interest. Of the three shoots emerging from the centrally located point of origin, one grows vertically. This, however, breaks off while still within the “Palaeozoic” circle. A second shoot, by contrast, extends to the upper right almost until it reaches the edge of the sheet, having begun to divide, while still within the “Palaeozoic” circle, into further shoots that themselves go on to do so. The inclusion of instances that might, in retrospect, be viewed as evolutionary “errors” alongside those that are, in every sense, to be termed a “success” may be seen as further evidence of Darwin’s endeavour to accommodate variety in all its manifestations. A further, even more compelling instance is to be found in the third shoot issuing from the point of origin in Darwin’s drawing. This initially runs towards the upper right, but very soon turns around in order to proceed in almost the opposite direction, although eventually proving prolific even within the first circle. While not a full return to its starting point (which might be an alternative way of viewing

1. The circle segments

23 _ Charles Darwin, Drawing of evolution (species transmutation) through a process of “natural selection”. Pencil on blue paper, 1851–57. Cambridge University Library, DAR 205.5: portfolio of loose sheets, fol.  184r. Reproduced with the permission of the Syndics of Cambridge University Library and of William Huxley Darwin

24 _ Charles Darwin, Drawing of evolution (species transmutation) through a process of “natural selection”. Pencil on blue paper, 1851–57. Cambridge University Library, DAR 205.5: portfolio of loose sheets, fol.  184v. Reproduced with the permission of the Syndics of Cambridge University Library and of William Huxley Darwin

the fate of the first shoot), this remarkable change of direction signals, nonetheless, an extremely rapid rate of mutation. While the two drawings described above might well be characterised as loosely “diagrammatic”, the third, drawn on the verso of Darwin’s blue sheet (Fig. 24), is rather distinct in this respect. Although inscribed, at its notional point of origin: “Parent of Rodents [deleted] Placentalia & Marsupalia”, this entity genuinely does resemble a drawn record of coral as encountered in the natural world, its “main” stem tending to the upper right, yet emitting, as it progresses, sequentially bi- and trifurcating shoots that, taken together, advance in every possible direction, their most distant termini forming an arc from upper left to lower right that follows the very faintly indicated circumference of a 30-degree segment of a circle. While Darwin, in this drawing, has eschewed the distinctive use of dotted lines as found in his first coralline sketch (Figs. 6, 7), one could not possibly claim that the form he has drawn resembles a tree.

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IV. From the circle drawings to the diagram (1851–1858)

All three drawn forms (Figs. 20, 23, 24), on account of their more or less “sprawling” character, may be said to resemble Darwin’s most important coralline sketch of 1837 (Fig. 14), albeit this is now combined with the concept of time, explicitly embraced through the use of a circular model. These were very promising advances, which might well have enabled Darwin to bring a new dimension to his efforts to find a way of representing a chronological process on a planar surface. Darwin was, however, constrained to call a sudden halt to his investigations and related thought-experiments when, in the summer of 1858, he was overcome by a not unjustified fear that he might soon no longer be able to claim priority in his research and its chief outcome: his success (as he now began to believe) in explaining species transmutation through a process of “natural selection”.

2. Darwin re-reads Wallace It would seem that Darwin had first paused during his work on these drawings when it came to his notice, in the mid-1850s, that conclusions almost uncannily similar to his own on “natural selection” as the engine of species transmutation were perhaps being reached by the naturalist Alfred Russel Wallace. This much younger man was at this point engaged in travelling around the Malay Archipelago in search of rare zoological specimens – above all, beetles and other insects, butterflies and birds – for provision, through a London agent, to the Natural History collections of both museums and independent scholars. Wallace had been publishing on his own research for some time and Darwin had already become aware of his work. In an essay of 1856, which appeared in the widely read Annals and Magazine of Natural History, Wallace (evidently inspired by Strickland) had arranged bird families within the order Fissirostres (sedentary birds) and those within the order Scansores (climbing birds) in very simple diagrams, intended in each case to convey a notion of relative “affinities” between the families included (Fig. 25).3 In doing so, however, he had eschewed Strickland’s cartographic iconography. The distances between the distinct groups, which according to Strickland, were the outcome of extinctions at earlier periods, were shown by Wallace in terms of relative positioning along a linear frame. In its abstraction, this recalled Lamarck’s own merging of 3  Wallace 1856, pp. 193–216; on the Fissirostres pp. 196–207, with diagram on p. 206; on the Scansores pp. 207–216, with diagram on page 215. “It is intended that the distances between the several names should show to some extent the relative amount of affinity existing between them; and the connecting lines show in what direction the affinities are supposed to lie. By referring to the diagram it will be seen that there are seven families placed close together, forming a central mass. Beyond the Trogons at some distance come the Goatsuckers and Swallows, while at the greatest possible distance from each other are the Hornbills and the Hummers, the former having a distant affinity to the Kingfishers, the latter to the Swallows”(p. 206). Cf. Brooks 1984, p. 120; on Darwin and Wallace: Canguilhem 1994, pp. 104–106; Browne 2002, pp. 14–42; Shermer 2002, pp. 88–90.

2. Darwin re-reads Wallace

25 _ Alfred Russel Wallace, Diagram of the Affinities of the Fissirostres, accompanying his article published in the September 1856 issue of the Annals and Magazine of Natural History

ladder and tree models (Fig. 5). As Wallace employed only a single, central vertical, the tree model here effectively recurred in a simpler, purer form. In an essay published in 1855 in the same journal, Wallace had in fact already invoked the notion of a symbolic oak tree. And his use of the arboreal model was there especially compelling because he had not introduced it as a solution to the problem of how the affinities between various groups (be they families, genera or species) might best be represented, but rather as a motif of the severe complications encountered in any attempt at overcoming this problem: “[There is] difficulty [in] arriving at a true classification, even in a small and perfect group; – in the actual state of nature it is almost impossible, the species being so numerous and the modifications of form and structure so varied, arising probably from the immense number of species which have served as antitypes for the existing species, and [have] thus produced a complicated branching of the lines of affinity, as intricate as the twigs of a gnarled oak or the vascular system of the human body”.4 4  Wallace 1855, pp. 184–196, here p. 187. Cf. Di Gregorio 1983, pp. 245–47; Brooks 1984, pp. 74–75; Shermer 2002, p. 85.

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IV. From the circle drawings to the diagram (1851–1858)

Only when, argued Wallace, it had become possible to grasp the complexities of the tree as a symbol of the species in their true chronological dimension would scholars be in a position to embark on classifying nature in its entirety: “Again, if we consider that we have only fragments of this vast system, the stem and main branches being represented by extinct species of which we have no knowledge, while a vast mass of limbs and boughs and minute twigs and scattered leaves is what we have to place in order, and determine the true position each originally occupied with regard to the others, the whole difficulty of the true Natural System of class­ ification becomes apparent to us”.5 Wallace here employed the image of the tree in order to present nature as a process in which many of the most important events had taken place in an unimaginably distant past epoch. This concept, deployed so powerfully by Wallace, was to prove immensely far-reaching. It was through reference to the image of a tree that he was moved to pose the question as to whether any particular aspect of change in the natural world could, unequivocally, be considered as pure “regression” or as pure “advance”. He evoked the image of a branch of powerful growth and positioned high up in a tree, that yet, as an outcome of unfavourable circumstances, breaks off, with the result that, far down the tree, another, much younger branch is additionally favoured in its own attempts to flourish. Occupying, however, a lower “rung” on the ladder of evolution, this second branch is possessed of the drive, but not of the strength and distinction, of the first branch. And yet, the change in circumstances, so injurious to the older and stronger branch, offers the younger, weaker branch a greater chance to thrive.6 Darwin had read the article of 1855 upon its appearance. It must have impressed him not only on account of the verbal eloquence with which Wallace employed the notion of a tree, as a compelling token of the difficulties of encompassing the species and their history in any form of model; but also, and above all, because he would surely have been taken aback to discover that the trunk and the main branches of Wallace’s notional tree so closely corresponded with how he had himself come to think of the petrified stems of a coral formation. If, in Wallace’s metaphor, the trunk and stronger branches equated to those extinct species of which it was thought no longer possible to obtain any direct knowledge, then this may well have had the effect of drawing Darwin’s own imagination (which had taken him from a tree model towards a coral model) back to the metaphor of the tree.

5 

Wallace 1855, p. 187. Cf.Brooks 1984, pp, 74–75; Shermer 2002, p. 85). “Thus every case of apparent retrogression may be in reality a progress, though an interrupted one: when some monarch of the forest loses a limb, it may be replaced by a feeble and sickly substitute” (Wallace 1855, p. 191). Cf. Brooks 1984, p. 76; Shermer 2002, pp. 85, 87. 6 

2. Darwin re-reads Wallace

At the same time, Darwin could have taken no pleasure in Wallace’s own use of metaphor: for the younger man, in employing the image of the upward striving tree and, moreover, identifying this as an oak, had thereby reintroduced that traditional hierarchy of evolutionary progress that the older man had himself abandoned. When, in 1857, Wallace had written to ask Darwin’s opinion of his 1855 article, Darwin had responded encouragingly enough, but had not refrained from adding: “I believe I go much further than you”.7 By now a number of Darwin’s colleagues had warned him that it was looking increasingly likely that Wallace would steal his thunder. This friendly advice prompted Darwin to look again at Wallace’s 1855 essay. The traces of this rereading are to be found in a number of Darwin’s marginalia (made in what appears to be a harder pencil than were his initial markings, and so leaving somewhat fainter lines), among which is a tiny, now barely visible drawing that comprises two broken lines roughly parallel to the edge of the printed text, one of them eventually trifurcating (Fig. 26). While ostensibly prompted by a passage in Wallace’s text distinguishing between “simple” and “diverging” lines of descent – the former to be represented by a “straight line” and the latter necessarily by a “forked, or manybranched line” – the form of Darwin’s annotation is astonishing: for, if turned upside down, it effectively creates a bridge spanning an interval of twenty years: back to 1837 and the first coralline sketch in his Notebook B (Figs. 6, 7).8

26 _ Charles Darwin, drawing in the margin of page 186 (on “straight” and “diverging” lines of descent) of article by Alfred Russel Wallace in his copy of the September 1855 issue of the Annals and Magazine of Natural History. Pencil, c.1857. Cambridge University Library, DAR. LIB.PER.U252. Reproduced with the permission of the Syndics of Cambridge University Library and of William Huxley Darwin 7  Letter of 22 December 1857 from Darwin to Alfred Russel Wallace, in: Darwin: Correspondence, Vol. VI (1990), no. 2193, pp. 514–15. The overall tone of Darwin’s letter is, however, remarkably encouraging, ending with enthusiastic comments on Wallace’s many years in the Malay Archipelago and his plans to stay on there for a while: “What a wonderful deal you will have seen […] I infinitely admire and honour your zeal and courage in the good cause of Natural Science”. 8  Darwin’s copy of the volume of the Annals with Wallace’s 1855 essay is now in Cambridge University Library, DAR.LIB.PER.U252.

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IV. From the circle drawings to the diagram (1851–1858)

Recognising a very real risk of missing his chance to establish his priority in formulating the explanation of species transmutation through a process of “natural selection”, Darwin now re-adopted the undeniably compelling tree model, and was indeed in due course to pass it off as his own invention. In fact, when summarising Wallace’s 1855 article for his own records (which he would have done, as was his habit, after a first reading), he had already noted on a sheet inserted in the back of the volume: “Every species has come into existence coincident in time and space with pre-existing species” (here quoting an approximation of Wallace’s own words), then added: “Uses my simile of tree”.9 Darwin’s comment here is vexing because the tree “simile”, or metaphor, which he had himself considered and then again rejected, could in no sense be regarded as his own invention. He had yet, moreover, to publish even a line establishing it as such. Finding himself in far more serious rivalry with Wallace than he had ever expected to be, he now revived the very way of envisaging evolution that he had overcome in the thought-laboratory of his notebook sketches and subsequent drawings. It was the risk of an unwanted competition with the younger man that prompted Darwin to put his misgivings aside and to “re-work” his own theory of evolution in terms of a tree model. He had recognised that, in view of the eloquence with which Wallace had established the “tree of life” as a symbol of perceptual and intellectual difficulty, it would not now be possible to arrive at an equally compelling alternative model for nature in its entirety. A letter that Darwin sent to his esteemed colleague Asa Gray, the pre-eminent American botanist, on 5th September 1857 marks the point by which he had resolved to pass off the tree metaphor as an original idea of his own. For along with this communication he enclosed a document (neatly copied out for him from his own manuscript) – “the briefest abstract of my notion on the means by which nature makes her species” – which purported to summarize the essence of his thinking to date, and which included the statement: “This, I believe, to be the origin of the classification or arrangement of all organic beings at all times. These always seem to branch and sub-branch like a tree from a common trunk; the flourishing twigs destroying the less vigorous – the dead and lost branches representing extinct genera and families”.10

  9 

For a detailed discussion of Darwin’s annotation and summary (albeit with no reference to his sketch of 1837), see Brooks 1984, pp. 244–48. 10  Letter of 5 September 1857 from Darwin to Asa Gray, in: Darwin: Correspondence, Vol. VI (1990), no. 2136, pp. 445–49 (enclosure: 447–49), here pp. 446 and 449. The manuscript from which the enclosure sent to Gray was copied is preserved: Cambridge University Library, DAR 6.51: “sketch of species theory sent to Asa Gray, 1857”. Characteristically, Darwin ended this communication to Gray with the avowal: “This sketch is most imperfect; but in so short a space I cannot make it better. Your imagination must fill up very wide blanks.–” Cf. Brooks 1984, p. 207.

2. Darwin re-reads Wallace

In the extensive manuscript on which Darwin was at work between 1856 and 1858, but in fact never completed (initially referring to it as his “big book on species”, though already telling Gray that it was to bear the title “Natural Selection”),11 he both adopted Wallace’s tree metaphor (albeit not in the role it had played in Wallace’s text) and reinforced this through his own arguments. He now wrote of the branches and twigs that may be understood to represent living species, and of dead branches that can be seen to represent those that are extinct. And he now claimed that it was only very rarely possible – in this he shared Wallace’s scepticism – to establish any detailed knowledge concerning long extinct organisms (on account of the incompleteness of the fossil record).12 Darwin recognised that the image of the tree – the profusion of its branches fallen to the ground, juxtaposed with the continually emerging new shoots, which are in turn engaged in drawing on those resources still required by the sappy, flowering branches – was a fitting image of nature ever re-creating itself out of a permanent state of catastrophe. With an eye on the writings of Wallace, Darwin now found that his understanding of evolution, which he had previously seen as manifest in the “sculptural” form of the dead stem of a coral formation, might be far more stirringly evoked and communicated in the form of the “battle painting” of a simultaneously dying and blossoming tree. And so, for the first time since his assertion, in his Notebook B, that “organized beings represent a tree”, he now returned to this model and, moreover, in emphatically macro-evolutionary terms: “[…] so by generations I believe it has been with the great Tree of Life, which fills the crust of the earth with fragments of its dead & broken branches, & covers with its ever living, ever diverging & marvellous ramifications, the face of the earth”.13 Wallace’s own use of the tree as a token of scepticism gave rise, in Darwin’s freshly provoked imagination, to the ravishing evocation of a tree that nature had brought forth as an all-encompassing symbol of evolution. In taking this new step, however, Darwin not only cancelled the critical distance between himself and Wallace; he also relinquished some of the notions that had once clearly most appealed to him. He abandoned his former intellectual allegiance to the anarchy of the circle, open on all sides, favouring instead the linear arrangement of time that Strickland had disdained as a fetishisation of the longing for order. From this point on Darwin would be constrained to enact his own embrace of self-contradiction, promoting an arboreal model about which he was, at heart, not entirely convinced in order to salvage his claim to priority.

11 

Letter of 5 September from Darwin to Asa Gray, in: Darwin: Correspondence, Vol. VI (1990), no. 2136, pp. 445–49, here 447: “I think it can be shown that there is [a] power at work, or Natural Selection (the title of my Book) […]”. 12  Darwin: “Natural Selection” (ed. 1975), p. 249. 13  Darwin: “Natural Selection” (ed. 1975), p. 250).

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IV. From the circle drawings to the diagram (1851–1858)

3. The precision of Darwin’s initial diagram The conflict also played itself out as a dichotomy between text and image because Darwin’s visualisation preserved traces of his residual reluctance to propose the “tree of life” as a model of species transmutation as he now understood it. But his scheme did not comprise a diagrammatic re-working of a verbal description of the tree model. It was, rather, an autonomous entity that dispensed with everything that was obviously mimetic of the natural world. This will be especially evident if one first considers the tree model devised by a German contemporary, the Heidelberg zoologist Heinrich Georg Bronn. Darwin was familiar with Bronn’s publications, and had indeed found there many an anticipation of his own ideas.14 Bronn’s own research, as Darwin came to realise, had sometimes brought the German “terrifyingly close” to the theory of evolution that Darwin was himself to present. Yet every time Bronn pondered more deeply on the significance of particular phenomena, he seemed to “de-fuse” the conclusions he might have drawn through encompassing these within the theory of recurrence to an original type, as this was to be expounded in his own publication of 1858, on the Entwicklungs-Gesetze der orga­ nischen Welt [Laws Governing Development in the Organic World].15 In his conclusion to this work, Bronn published the simplified image of a leafless tree, the branches of which were intended to signify the various living organisms in the chronological sequence of their emergence (Fig. 27). According to Bronn,

27 _ Heinrich Georg Bronn, Tree model of the emergence of living organisms. Illustration in his 1858 volume Entwicklungs-Gesetze der organischen Welt 14  15 

Junker 1990, pp. 187–88. Junker 1990, pp. 186, 188.

3. The precision of Darwin’s initial diagram

this tree showed the development of life within an environmental context that itself remained fixed as initially established. Each of the living organisms in Bronn’s arboreal diagram evolves in a manner not unlike that envisaged in the transformative scheme proposed by Lamarck; and, in so doing, it continually threatens to impinge upon the domains of others. But not one of these ever transgresses its own boundaries – as would seem to be an all but inevitable outcome of the situation evoked in some of Darwin’s own drawings (Fig. 20) – merely filling out the space available to it. In Bronn’s diagram the evolution of living organisms is to be read from the bottom up: from the invertebrate animals to humankind. Here, an elevated level of development in a lower life form may, on occasion, attain a greater degree of “perfection” than is to be found in the original phase of a higher life form. Thus, for example, on the left a twig marked “d”, growing from the branch labelled “B”, has attained a higher degree of development than has the twig marked “c”, growing from the branch on the right labelled “C”, even though the latter itself grows from a higher point on the trunk. And the vertically tending, but then eventually terminating, twigs ultimately show that each type of organism emerges, endures for a certain period of time, and then departs the scene. Taken together, these aspects of Bronn’s model nonetheless give an initial visual impression of living entities that, in their evolutionary zeal, might well at some point overstep the boundaries imputed to them in Bronn’s text. In short, Bronn’s image as a whole strikes the eye as far more adventurous than are his words.16 Darwin’s diagrams are, however, not comparable with Bronn’s model – not least on account of the mimetic character of the latter. Darwin’s initial version of the diagram eventually published in 1859 in his volume on the Origin of Species (Fig. 33) was devised in around 1857 (Fig. 29), but was itself first published, in a reconstructed form, only in 1975 (Fig. 28). This initial version reveals, in particular, Darwin’s attempts to achieve the sort of schematic clarity that would be likely to provide the reader with a rapid overview of a phenomenon of enormous complexity and a process of unimaginably long duration. Darwin intended that this diagram be published as an essential component of his argument, and that it should itself repeatedly engage the attention of the reader. As he was to note, astutely preempting the needs and limitations of those who might in future peruse his book: “the complex action of these several principles, namely, natural selection, divergence & extinction, may be best, yet imperfectly, illustrated by the following diagram, [to be] printed on a folded sheet for convenience of reference”.17 The single printed diagram of 1859, positioned, as Darwin had proposed, as a fold-out sheet bound in with the text, comprises, in essence, a merging of the initial, drawn diagrams that Darwin labelled “I” and “II” (Fig. 29), albeit with the initial 16  17 

Bronn 1858, pp. 481–82; cf. Uschmann 1967, p. 14. Darwin: “Natural Selection” (ed.1975), p. 238.

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IV. From the circle drawings to the diagram (1851–1858)

28 _ Reconstruction of Darwin’s initial diagram of evolution (species transmutation) of c. 1857 (see Fig. 29), published in “Natural Selection” (ed. 1975)

3. The precision of Darwin’s initial diagram

29 _ Charles Darwin, Initial diagram of evolution (species transmutation) through a process of “natural selection”. Pencil, pen and ink, body colour, printed letters, adhesive, c. 1857. Cambridge University Library, DAR 10.2: manuscript of “Natural Selection” chapter six, after fol.  26r. Reproduced with the permission of the Syndics of Cambridge University Library and of William Huxley Darwin

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IV. From the circle drawings to the diagram (1851–1858)

top-down reading replaced by a bottom-up arrangement. In the drawn diagram the aligned parent-species within a single genus are represented by large capital letters.18 Darwin then mentally pursues (Fig. 29, upper section) how each of these parent-species evolves over thousands, even millions of generations, right down to a nominal “present day”. By contrast with those parent-species labelled “B” to “L”, which persist for a while unaltered and then become extinct, parent-species “A”, through its distinctive fan-like formations, brings forth marked varieties. But, of these varieties, and in due course sub-varieties, only a few prove capable of surviving. And so, at the end of ten units of generation (every one of which represents a vast number of real-time generations), only those marked “a”, “h” and “l” are found to have survived. In the case of parent-species “M”, on the other hand, a less vigorous form of divergence is found to occur, repeatedly producing only the varieties “z” and “m”. Darwin was here, perhaps, influenced by Wallace’s theory of the frequent existence of moribund “antitypes”; and in the present case it is evident that, by the end of the sequence of descent shown here, only the “m” line endures. Darwin’s diagram II (Fig. 29, central section) abstracts this sequence of developments to the successful lines of descent and registers in more detail alternative developments, in addition to further evidence of the capacities of the parent-species. Diagram III (Fig. 29, penultimate line) then shows the outcome after ten chro­nological units of evolution; and diagram IV (Fig. 29, last line) reveals that, after the passage of a further vast and indeterminate span of time, six new subvarieties have evolved from the two earlier varieties of the parent-species “A”; that parent-species “E” and “F” have persisted unaltered; and that parent-species “M” has now given rise to three new varieties. All the other species have become extinct. It is immediately clear that Darwin’s diagrams I and II differ from Bronn’s mimetic model in that they are intended to be read, as was the case with Lamarck’s diagram (Fig. 5), from the top down. It is, moreover, of no small significance that Darwin makes no attempt to suggest branch-like forms with his columns of lowercase letters and superscript numerals; and that he makes use of dotted lines, as he had done, in a manner perhaps indebted to Lamarck, in his first coralline sketch of 1837 (Figs. 6, 7) and in a detail of one of his drawings of the 1850s (Fig. 22). An element fundamental to Darwin’s diagram – its downward pointing bifurcations – also suggestively reiterates the form he had drawn in the margin of Wallace’s 1855 article (Fig. 26). The introductory description that Darwin intended should accompany his diagram emphasises the degree of uncertainty within it. This was quite a contrast with the suggestive character of the tree model devised by Bronn (Fig. 27): “This diagram will show the manner, in which I believe species descend from each other & therefore shall be explained in detail: it will, also, clearly show several points of 18 

Darwin: “Natural Selection” (ed. 1975), p. 239.

3. The precision of Darwin’s initial diagram

doubt and difficulty”.19 In his text Darwin also entertains numerous possible explanations as to how species may come to remain stable or alter at different rates, repeatedly returning to an insistence that, without the diagram, it would be impossible to gain any conception of the overall process of “natural selection” and that, on this account, it should also be borne in mind that any such diagram must rely on a substantial degree of schematic abbreviation. He thus characterises parentspecies “M”, as shown in the diagram, as an ideal from which conceptual generalisations should not be drawn: “I do not suppose the process generally to have been so simple as represented under M, where a simple variety m1—10 in each stage of descent has been naturally selected”.20 Notwithstanding his concern for clarity of exposition, Darwin goes quite some way towards disturbing and confusing the harmonic symmetry of his diagram. This endeavour begins, indeed, with his introductory explanation regarding the positioning of the letters identifying parent-species “A” to “M”: “The unequal distance of [= between] the letters may represent the ordinary way in which the species, even when as in this imaginary case all are closely related together, yet stand unequally related in little sub-groups”.21 In view of the prominence of this advice to the reader, it is most regrettable that, in the version of Darwin’s original diagram included (as a putative exact rendering of this document) in the otherwise most valuable 1975 publication of the “Natural Selection” manuscript (Fig. 28), precisely this aspect is misrepresented. As the capital letters there are positioned with always exactly the same space between one and the next – as if they were troops mustered for inspection – they are deprived of that element of irregularity by which Darwin set such store. All the greater, then, is the surprise that awaits the researcher fortunate enough to be able to inspect the original document,22 who will be all but “ambushed” by a large, pale blue, very faintly lined sheet of paper, which at roughly 30 by 40 cm has the dimensions of a poster (Fig. 29). Here, each of the “headline” letters in diagrams I through IV takes the form of a large capital, first printed, then cut out, then glued on to the sheet, then emphasised through the addition of a full-stop drawn in firmly with pen and ink. Over time the glue has assumed a brownish tinge, which now lends this parade of capitals an unruly, almost shabby air – although this too, ironically, complements Darwin’s endeavour to avoid, at all costs, the regularity of a conventional diagram. It would have been in keeping with this concern that Darwin, in diagram I, also drew in with pencil, rather sketchily and irregularly, the

19 

Darwin: “Natural Selection” (ed. 1975), pp. 238–39. Darwin: “Natural Selection” (ed. 1975), p. 242. 21  Darwin: “Natural Selection” (ed. 1975), p. 239. 22  Cambridge University Library, DAR 10.2: “Natural Selection”, chapter six, unnumbered sheet after fol. 26r. 20 

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IV. From the circle drawings to the diagram (1851–1858)

30 _ Detail (uppermost section) of Fig. 29

letters “N” and “O” next to the “M” that is the last of the affixed printed letters. The irregularity in the row of “headline” letters required by Darwin was itself achieved through his cutting away the paper of the printed sheet lying between the contiguous letters “C” and “D”, “D” and “E”, “F” and “G”, or “L” and “M”, so as to be able to vary the intervening distances, and so determine the “rhythm” of the sequence as a whole. In the published reconstruction of 1975 (Fig. 28) one finds nothing of this. This approach is also true of the subtlety Darwin brought to inscribing the further details in the initial document (Fig. 29). In diagram I, for example, one finds that from the capital letters “A” and “M” there extend vertical dotted lines, and from capital letter “A” also diagonal dotted lines, in all cases identified by lowercase letters and superscript numerals (Fig. 30). In the case of “A” the uppermost groups of dots are marked in with pen and ink. Further down the sheet only the central vertical is so marked, while the diagonals are faintly drawn in, as continuous lines, with pencil. In addition to these markings, there are traces of pencil (often so faint as to be invisible in reproduction) that are indicative of under-drawing, corrections and additions. From the point marked i6, for example, there extends a dotted bifurcation to the level of the line running between i7 and ll7 that is barely perceptible even with the aid of a magnifying glass. And from z7 to points lying to the left of m10 there extends an effective triangle of pencil markings, which Darwin at no point reinforces for emphasis with pen and ink (Fig. 31). And so this sprawls, like a shadow, along that z-line, which – unlike its m-line pendant – ultimately proves able to sustain no varieties. This aura of irregularity is the defining characteristic of Darwin’s initial diagram. It also very movingly reveals what sensitivity this naturalist, albeit so ungifted as a draughtsman, brought to the task of visualisation. Affixed at the upper right, finally, were to be found Darwin’s instructions to the compositor (Fig. 32), these too referring to the intentionally irregular distances

3. The precision of Darwin’s initial diagram

31 _ Detail (lower right section) of Fig. 29

between each of the “headline” capital letters, and going into great detail regarding other aspects of how the diagram should be type-set: “Compositor: To be printed on separate page to be folded out and so all exposed. Attend to distance of capital letters from each other: the letters had better be smaller: attend to chains of dots and hyphens. The numbers to small letters to be the very smallest possible. The capital and other Letters in each diagram to match exactly in position. – I hope the 4 Diagrams will go in length of page. –” The care with which Darwin glued, inscribed, drew and explained his original diagram – his model of the evolution of species through the process of “natural selection” – compels admiration. It is thus all the more disheartening to contemplate how insensitively the printed version of 1975 (Fig. 28) falls short of this model.

32 _ Darwin’s directions to the compositor (affixed to upper right of Fig. 29) on the printing of the diagram. Pen and ink on paper

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IV. From the circle drawings to the diagram (1851–1858)

It effaces every linear differentiation, with the result that it is no longer clear which markings were, often hesitantly, in pencil and which, more definitively, in pen and ink. Nor is there any differentiation between distinct types of drawn line, those in the initial version being by no means always straight. A “talented” designer has here eradicated the traces of Darwin’s unruly, but also tentative, drawing hand. And, along with this, everything that so enlivens Darwin’s diagram has been removed.

V. The fold-out diagr am in the Origin of Species (1859)

1. Form Darwin’s endeavours of 1856–58 to concentrate the fruit of over two decades of investigation and reflection into a single, long text on “natural selection” was rather more suddenly interrupted when, in the summer of 1858, he received a package sent by Alfred Russel Wallace. This contained the manuscript of a further essay (written, like those of 1855 and 1856, during his travels around the Malay Archipelago), and the request that Darwin, if he thought the text had merit, be so kind as to pass it on to the eminent geologist Charles Lyell (who, as Darwin had himself informed Wallace, had greatly admired the 1855 text). To Darwin’s astonishment, Wallace’s new manuscript contained no less than the effective essence of Darwin’s own conclusions: “I never saw a more striking coincidence”, he wrote to Lyell, adding in a spirit of resignation: “So all my originality, whatever it may amount to, will be smashed”.1 Lyell, however, in league with the botanist Joseph Hooker, rallied in Darwin’s support, urging that he hasten to devise a publishable summary of the research, discoveries and theory recounted in his long manuscript, lest Wallace soon come to be declared the first to have fathomed the workings of species transmutation through the process that Darwin now termed “natural selection”. Having secured Darwin’s still rather half-hearted agreement to this proposal, they then arranged, at

1  Letter of 18 June 1858 from Darwin to Charles Lyell, in: Darwin: Correspondence, Vol. VII (1991), no. 2285, pp. 107–08, here, 107. The arrival of the package from Wallace has usually been assumed to pre-date by only a day or so the above-mentioned communication. This assumption has, however, been questioned by several scholars, most notably Brooks 1984, pp. 248–57. He argues (on the basis of detailed research into postal communications from the Malay Archipelago at the period in question) that the package may well have arrived up to a month earlier, regarding “May 17–18” as “the most likely time” for it to “have reached Darwin’s hands” (p. 256), thus giving the recipient sufficient opportunity to incorporate aspects of Wallace’s argument into his manuscript before writing to Lyell. These views have not, however, found general assent among scholars. On this matter, see: Darwin: Correspondence, Vol. VII (1991), Introduction, pp. xvii–xvii.

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V. The fold-out diagram in the Origin of Species (1859)

very short notice, for a joint, Darwin-Wallace presentation to be added to the agenda for the 1st July meeting of the Linnean Society in London, under the single title: “On the tendency of species to form varieties, and on the perpetuation of varieties and species by natural means of selection”. This comprised readings, in turn, of part of Darwin’s account of his theories as committed to paper in 1844; of an extract from the manuscript from which had been copied the outline sent in 1857 to Asa Gray; and of the full text of Wallace’s new essay.2 Neither Darwin nor Wallace was in attendance. (Darwin’s infant son, who had recently succumbed to scarlet fever, died on 28th June; and Wallace was not in fact to return to England until spring 1862.) But, between the summer of 1858 and the spring of 1859, Darwin devoted all his time and energy to devising an abbreviated version – in his own words, an “abstract” – of his incomplete 1856–58 manuscript, omitting all the data and information on sources originally supplied in footnotes. These he now proposed to include in a later, fuller version of the work – though in fact he never did so. Proofreading occupied the summer and early autumn of 1859, all but exhausting the author. The resulting, shorter text – its first printed version, nonetheless, running to no fewer than 490 pages – was the Origin of Species as published in London on 24th November 1859. Tellingly, however, for all Darwin’s ferocious pruning, the planned fold-out sheet was not omitted. It was bound, between pages 116 and 117, in the last third of his longest and most important chapter, now chapter four (formerly chapter six): “Natural Selection”. And the diagram it bore (Fig. 33) was to become one of the most influential images ever brought forth in the realm of the Natural Sciences.3 The printed version of the diagram retained the same basic elements as were to be found in the model supplied within Darwin’s manuscript (Fig. 29). The parentspecies were indicated with capital letters (albeit with the initial sequence “A” to “M” now reduced to “A” to “L”); their evolution over time was charted through both vertical and, chiefly, diagonal dotted lines; and the varying degrees of differentiation between the parent-species were also signalled: while the groups “A” to “D” and “G” to “L” were set appreciably apart from the pair “E” and “F”, these two series also exhibited slight, but still significant, variations in the distance between one letter and its neighbour(s).

2 

Wallace’s essay was, accordingly published far more promptly than he could have dared hope when sending it to England from the other side of the world. See: “On the Tendency of Varieties to Depart Indefinitely from the Original Type”, Journal of the Proceedings of the Linnean Society: Zoology, III/9 (20 August 1858), pp 53–62. The author subsequently ensured his text of a much broader readership through including it, essentially unaltered, as the second of ten, largely pre-published articles (and directly preceded by that of 1855), in the volume Contributions to the Theory of Natural Selection, London 1870, pp. 26–44. 3  Gruber 1988, p. 132.

1. Form

33 _ Charles Darwin, Diagram of “natural selection” published as a fold-out sheet bound into chapter four of the first, 1859 edition of the Origin of Species

In view of this fidelity to Darwin’s initial diagram, the instances where the printed version differed are all the more striking. There is no record as to why these changes were made; but it would be fair to assume that they derived from Darwin’s further reflections, perhaps set in train by, but going far beyond, practical problems encountered with the arrangement as at first envisaged. Comments on the printed diagram made in Darwin’s correspondence with his publisher, John Murray, between April and June 1859, suggest that he had by that point recognised that the fold-out sheet bearing it might be markedly elongated in form (thus requiring a vertical fold or two), but should not be of such a height as to necessitate, in addition, a horizontal fold.4

4  See Darwin’s letters to Murray of 2 April, 14 May, 31 May and 14 June 1859, in: Darwin: Correspondence, Vol. VII (1991), respectively no. 2446, pp. 277–78; no. 2462, pp. 298–99; no. 2465, pp. 300–01; and no. 2469, p. 303. Although Darwin had specified that his new hand-drawn diagram (enclosed in the letter of 31 May) should be engraved, Murray in fact commissioned a lithographic version from the well-established medical and scientific illustrator William West. As had been requested, a proof of the diagram was promptly sent to Darwin, who on 14 June declared that the version provided by West would “do very well”.

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V. The fold-out diagram in the Origin of Species (1859)

In place of Darwin’s initial four separate (albeit vertically stacked) diagrams, a single diagram covered the entire fold-out sheet. The most significant departure, however, was that the printed version was intended to be read not from the top down, but from the bottom up. Also very telling is the further differentiation now introduced into the rendering of lines. The eleven lines that descend from the row of capital letters (and which had no equivalent in the initial diagram) are not dotted, but broken, that is to say made up of a sequence of very short dashes. (A hint of this is already to be found in a detail of one of the circle segment drawings: Fig. 22. It recurs in the form of Darwin’s marginal annotation to Wallace’s 1855 essay: Fig. 25.) These broken lines represent the vast, indeterminate span of past time, during which the parent-species may be assumed to have evolved from a common ancestor. This is intended to suggest to the reader that the element of separation within the groups of letters “A” to “D” and “G” to “L” would be maintained, albeit gradually diminishing, until the lines descending from each parent-species reach their shared origin, and that the resulting merged lines would eventually meet the likewise gradually merging lines descending from parent-species “E” and “F”. It is for this reason that all these lines are shown so as to make clear that they eventually converge (that is to say, had had a common origin), albeit in a process so infinitely slow as to be all but unimaginable. In the uppermost section of the printed diagram a corresponding series of broken lines ascends through the horizontal bands marked XI to XIV, each of these representing an interval of future time, during which any descendants of the parent-species still in existence in horizontal band X will themselves further evolve. These descendants, here considered (for the sake of expositional clarity) as “varieties” / “sub-varieties”, may well be sufficiently diversified and distinct – as is indeed implicit in their persistence – to qualify as new species in their own right. The largest, central section of the diagram, which lies between horizontal bands I and X, represents a considerably shorter span of time than that traversed by the broken lines in the sections found below and above it. Even though the dotted lines must be imagined as making their way through chronological intervals that each comprise a thousand generations, theirs is nonetheless a far shorter journey than that undertaken by the broken lines. In the zone traversed by the dotted lines Darwin effectively offers his readers a segment of his entire diagram as viewed through a magnifying glass, with the aid of which they are able to detect the emergence of significant variations among the descendants of the parent-species. The printed version of the diagram in its entirety thus presents a simultaneous record of the view obtained from diverse chronological perspectives. The horizontal bands I through X show how the two dominant parent-species “A” and “I” give rise to ever new varieties, here recorded in the form of dotted lines of different lengths ascending diagonally. On the whole, it is the outer diagonal dotted lines that emerge as victors in the struggle for survival, while the others (in

1. Form

addition to the occasional vertical line), which simply terminate, represent varieties that relatively soon become extinct. Each of the bands I through X stands for a period of time, after which significant varieties of the parent-species “A” and “I” have emerged. By the advent of horizontal band X, parent-species “A” has given rise to three significant varieties (here marked a10, f10, and m10), while parent-species “I” has given rise to two significant varieties (w10 and z10), but parent-species “E” and “F” have persisted without giving rise to varieties that, in this context, are deemed significant (thus re-appearing at horizontal band X as, respectively, E10 and F10, the former apparently terminating at that point). In the radically simplified upper register of Darwin’s diagram, where the dotted lines are again down-graded to the thicker strokes of the broken lines, it is apparent that, alongside the persisting stable species “F”, all that will survive are further varieties, or sub-varieties, of the two dominant parent- species “A” and “I”. After the passage of the generations in existence during the time represented by the upper horizontal bands XI through XIV only fourteen further varieties, or sub-varieties, are to be found (eight ultimately deriving from parent-species “A”, six from parentspecies “I”), these co-existing with the persisting, essentially undiversified species “F”. Every other variety / sub-variety to have emerged at any point during the entirety of the period represented by the diagram will now be extinct. In rendering the persistence, variation and extinction of the species comprehensible “at a glance”, Darwin’s diagram, with its graphic visualisation of both magnitudes and processes, offers his readers an especially eloquent image. Through his accompanying commentary (its main segment occupying pages 116 to 126 of chapter four in the first, 1859 edition), Darwin sought to explain the interplay of the diverse graphic elements with both the letters and the numerals. He was subsequently to observe that, if the lineage of an ancient family could not be rendered even minimally comprehensible without resort to such a diagram, then this was even truer of the natural families of the species, the evolution of which would be impossible to grasp without such visual assistance and, even so, remained difficult to comprehend.5 It was, moreover, only in this diagram (and nowhere else in his published work) that Darwin sought to provide an account of nature in its entirety. Precisely because the endless variability to be found in nature evaded human comprehension in terms of a recognisable “system”, it must needs be grasped through the “description” afforded by an image. And herein lies the fundamental methodological significance of Darwin’s diagram of “natural selection” as the engine of species transmutation, and thus of evolution.

5 

Darwin: Origin of Species (1859), p. 431.

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V. The fold-out diagram in the Origin of Species (1859)

2. Relation to the text In as far as Darwin’s published diagram did rather more than demonstrate a mathematical truth – what he himself terms “the high geometrical powers of increase of each species” – 6 one is moved to enquire further into precisely what, in its capacity as a “description”, it was a “description” of. Commentators have as yet barely touched on this issue,7 above all because the published diagram, whenever it has been considered in this light, has invariably been assumed to be mimetic of the branching and emphatically upward directionality of a tree. While the dotted lines that ascend diagonally from the parent-species “A” and “I” do indeed evoke the impression of organic branching, this resembles not so much the branching characteristic of a tree as that of a shrub or a bush. This in turn prompts one to wonder whether Darwin really intended his diagram to serve as a visual rendering of the “tree of life” that features in the printed text, as it had in the text of his manuscript of 1856–58, or now wished the diagram and the text that accompanied it to stand in effective “competition” with each other. In the manuscript – Darwin’s text in its fullest form – this was evidently the case. Here, Darwin had evoked the model of a tree with a good deal of eloquence, only to avoid any hint of an arboreal structure in his initial, four-part diagram. And, while the evolution of parent-species “A” in diagram I (Fig. 29, upper left) could certainly be interpreted as a branching structure, the fact that this is a downward, not upward, ramification goes quite some way towards weakening such an association. It is, then, all the more telling that Darwin, in his description of the printed diagram (a diagram that does, after all, adopt an upward directionality) appears almost expressly to avoid employing the metaphor of a tree. Not in a single word does he here refer (as had Wallace, and so vividly, in his 1855 essay) to “leaves” or to “twigs” or to a particular species of tree. He makes use only of the much more neutral term “branches”, and then without any direct reference to an object in nature, but rather to the action of branching, understood geometrically. Darwin in fact here writes of his own diagram more geometrico – in terms, that is to say, of his particular use of lines (be they dotted or broken), and of letters and numerals. Indeed, throughout the rest of the volume, with only two exceptions8, Darwin’s use 6 

Darwin: Origin of Species (1859), p. 127. As far as I know, only Pörksen 1998, pp. 328–30 has so far posed this question. 8  In chapter two (Darwin: Origin of Species (1859), p. 46) the insect genus Coccus is characterised by a principal nervous system that “may almost be compared to the irregular branching of the stem of a tree”. And in chapter six (p. 187), in a discussion of the Articulata in general, we learn of features characteristic of the earlier organs giving way, over time, to more complex forms, a process there compared to “numerous gradations of structure, branching off in two fundamentally different lines”. 7 

2. Relation to the text

of the term “branching” is intended to refer not to the corresponding parts of a real tree, but to the geometrical notion of branching lines. At one notable point, in chapter nine, “On the Imperfection of the Geological Record”, he even employs a traditional turn of phrase to evoke the (in fact, his own) theoretical notion of the past and present species of the same genus being connected not in the manner of the constituent parts of a tree, but in “one long and branching chain of life”.9 Whenever Darwin, elsewhere in the text of the first edition of his volume, mentions the diagram featured in his chapter on “natural selection”, there is no indication at all that he is thinking of this in terms of a tree model. The fact that this identification has, nonetheless, appeared so persuasive is perhaps to be explained by the impact on many readers of the fervour with which Darwin, towards the end of the chapter, evokes the image of a tree in nature, complete with trunk, twigs and leaves, as a model of evolution understood as a struggle for life: “The affinities of all beings of the same class have sometimes been represented by a great tree. I believe this simile largely speaks the truth. The green and budding twigs may represent existing species; and those produced during each former year may represent the long succession of extinct species. At each period of growth all the growing twigs have tried to branch out on all sides, and to overtop and kill the surrounding twigs and branches, in the same manner as species and groups of species have tried to overmaster other species in the great battle for life”. Darwin goes on to emphasise that, while it is the general rule that “[of] the species which lived during long-past geological periods, very few now have living and modified descendants”, or can be encountered only “in a fossil state”, it may occasionally happen that a form unlikely to survive, here symbolised by “a thin straggling branch springing from a fork low down in a tree”, may be so favoured “by some [environmental] chance” that it is enabled to persist, thus for a while co-existing with more vigorous and better adapted forms.10 Darwin concludes by reiterating, word for word, a passage from his original manuscript, speaking of “the great Tree of Life, which fills with its dead and broken branches the crust of the earth, and covers the surface with its ever branching and beautiful ramifications”.11 By comparison with the structure of Darwin’s published diagram, this image of a tree offers a chronological encapsulation of the dynamics of the struggle for life as a drama of budding, growth, the triumph of supplanting

  9 

Darwin: Origin of Species (1859), p. 301. Darwin, Origin of Species (1859), p. 129. 11  Darwin, Origin of Species (1859), p. 120. Darwin included the metaphor of the “tree of life” in only one other page of his volume, and in that case rather in passing. In chapter nine, “On the Imperfection of the Geological Record” (p. 317), he characterises the gradual emergence of groups of species as the “branching of a great tree from a single stem”. 10 

67

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V. The fold-out diagram in the Origin of Species (1859)

rivals, the grimly ineluctable fact of being in turn supplanted by other rivals: a concentration of directionality and recurrence.12 The high-flown language with which Darwin, at the end of his chapter on “natural selection”, seeks to persuade his readers to think in terms of a tree, even while failing to anticipate this passage in the preceding explanation of his diagram, confronts these readers with a discrepancy that cries out for explanation. One reason that has been advanced is that Darwin’s closing metaphor of the “tree of life” was intended to refer to evolutionary processes from the macro-biological point of view, while his diagram was intended as a simplified account of these processes viewed in micro-evolutionary terms.13 Such an explanation would, however, be contradicted by the fact that both the dotted and the broken lines found in the diagram themselves evince macro-evolutionary characteristics. Far from serving to explain the functioning of a minute detail within the “great scheme of things”, Darwin’s diagram represents processes invincible in their power, of barely imaginable duration, and valid for nature in its entirety. Darwin does not engage in any detail with the metaphor of the tree once he has embarked on his description of the diagram. Conversely, he wastes not a word on the diagram once he has engaged more deeply with the metaphor of the tree. It seems more than likely that this puzzling “incoherence” is to be explained not by reference to the text itself, but to a crucial aspect of the context in which it was produced: that inner conflict into which Darwin was thrown when he resolved – in reaction to a re-reading of Wallace’s 1855 essay – that the “tree of life” was, after all, the best model for envisaging, and helping others envisage, the process of evolution as he had himself come to understand this.

3. A “coral model” for evolution Darwin’s textual evocation of a metaphorical tree does not stand in relation to his published diagram (Fig. 33) as would a concept to an illustration. And yet, the formal arrangement of the sequentially emerging varieties of parent- species “A” and “I” is not itself without an aspect that does insinuatingly mimic nature. This is above all true of the recurrent fan-like forms made up of several abbreviated dotted lines. These equate to those varieties and sub-varieties preserved only in fossil form. In the image of the tree, as invoked at the end of Darwin’s chapter on “natural selection”, he found a metaphor for extinct species in the branches that had broken off from the tree and lay around it on the ground. In the diagram, by contrast, the extinct varieties and sub-varieties, which have, by definition, been defeated in the struggle for survival, are not supine, but emphatically upright: still raising their 12  13 

Pörksen 1998, p. 329. Uschmann 1967, p. 17.

3. A “coral model” for evolution

34 _ Detail of lower left of Fig. 33, showing pattern of dotted lines extending from parentspecies “A” and from variety a1

arms aloft even in the rigidity of death. At the lower left of the printed diagram, for example (Fig. 34), this is the case with four of the six dotted lines extending from parent-species “A”, and with four of the five extending from variety a1. In the stark difference between these two approaches to acknowledging the phenomenon of extinction is to be found the reason why the diagram remains fundamentally incompatible with the tree metaphor. These two expositional devices differ from each other in terms of structural logic. It is doubtless for this reason that Darwin avoided linking them.14 The widespread interpretation of Darwin’s diagram as an abstract rendering of a tree derives from a misunderstanding. In reality, the diagram is much closer to functioning as if based on the model supplied by coral, its reiterated motif of “arms raised aloft” corresponding far less persuasively to the branches of a tree (let alone those already detached from the tree and fallen to the ground) than to the petrified limbs that are coral’s own fossil form. If both of the fundamental graphic elements shared by the published diagram (Fig. 33) and the diagram that accompanied Darwin’s manuscript (Fig. 29) – the use of dotted lines and of bi- and trifurcating diagonals – may be found to derive ultimately from his earliest coralline sketch (Figs. 6, 7), this is by no means a coincidence. It is, rather, an outcome of the same logic that underpins the crucial presence of dotted lines in one of the circle segment drawings (Figs. 20, 22). In seeking real-world examples from which Darwin’s “schematic coral” may have derived (though clearly more than one instance would have contributed, in some degree, to this process), one could do worse than recall the specimen that Darwin had found in January 1834 on the coast of Patagonia (Fig. 1), even while this is in fact a coralline alga, and not, as Darwin had at the time believed, a true coral. If now re-appraised with Darwin’s later work in mind, this organism may be seen to evince a striking similarity with the central section of the right half of the published diagram (Fig. 33): that part relating to the varieties of parent species “I” 14  As has been argued (Pörksen 1998, p. 329), Darwin’s stirring verbal evocation of a “tree of life”, which seems to call out for a corresponding visual model, in itself hampers the reader’s capacity for visualisation because it links the recurring cycle of the seasons with the linear progression of the history of organic life on earth.

69

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V. The fold-out diagram in the Origin of Species (1859)

35 _ Detail of centre right of Fig. 33, showing pattern of dotted lines representing descent from parent-species “I”

(Fig. 35). The similarity is fully evident, however, in the lateral reversal of the form in which the specimen has been mounted, and so preserved. And, if one superimposes the reversed and silhouetted form of this organism on the aforementioned section of the diagram (Fig. 36), one finds that the “fit” is far more precise than would have resulted from a looser sort of affinity. The necessarily always straight segments of dotted line in the diagram cannot of course fully coincide with the rounded forms of the aligned “bosses” of the coralline alga, which they shadow in loosely parallel tracks. (And the short, non-bifurcating isolated limb visible in Fig. 1 is here omitted.) But it would, for example, be a remarkable coincidence if the angle formed by the meeting, at z4, of the z and u lines of descent or the slight rightward shift of the line z6 to z8 had been plotted by Darwin without any recollection of the specimen he had discovered two decades earlier. This seems all the more so in as far as just such a recollection would appear to have accounted for that very form Darwin had tended to envisage, from the time of his first Notebook B sketches in 1837 (Fig. 6), in seeking to arrive at a fitting representation of species transmutation through a process of “natural selection”. A final piece of circumstantial evidence is afforded by coincidence in terms of scale: the maximum height of Darwin’s specimen, measured from the base of its stem, is 6.3. cm – precisely that of the corresponding segment of his diagram as it was printed in 1859 (Fig. 33). The illustrated superimposition (Fig. 36), in which both elements are enlarged for greater clarity, shows the one emerging as an informative “abstraction” of the other. It is no longer possible to establish precisely how Darwin arrived at the abstraction of his Patagonian specimen that is apparently to be found in his published diagram. He may have referred to a drawing he had himself made of it earlier, observing it from the side now invisible to us (and thus reversing the left-right presentation we now know), transferring this at a much later date to a preparatory

3. A “coral model” for evolution

36 _ Superimposition of silhouette of Darwin’s Patagonian specimen (Fig. 1, here reversed) on the pattern of dotted lines shown in Fig. 35

form of his printed diagram. It seems highly improbable that Darwin would have relinquished any specimen he had collected – notwithstanding the invariably large number of these – before he had made some drawn record of it, at the very least for his own future reference. And a simple outline drawing would have been within the capabilities even of an exceptionally weak draughtsman. Although Darwin was well aware of his own inadequacy in this respect, there are some indications that he nonetheless persisted in his endeavours to capture on paper that which he had collected and that which he had observed. He himself reported this with regard to specimens amassed during the H.M.S. Beagle expedi­ tion;15 and it is implicit, at a later point in his career, in his comments regarding small corals and similar organisms. In a letter sent in late 1846 to the physiologist William Benjamin Carpenter, Darwin requested the address of the draughtsman 15 

Darwin: Autobiography (ed. 1958), pp. 77–78.

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V. The fold-out diagram in the Origin of Species (1859)

this colleague had recently employed, and went on to describe the items he himself wished to have recorded: “The objects which I want chiefly drawn are minute corallines, & minute articulata & molluscata & their various organs; please say whether you think your artist would do such things well”.16 It would seem that Carpenter replied promptly, for Darwin was indeed able to engage the services of the draughts­ man in question, who was Samuel William Leonard. And in December of the same year Darwin wrote to his friend Joseph Hooker of his satisfaction with, and keen engagement in, Leonard’s work: “When I was drawing with Leonard, I was so delighted with the appearance of the objects, especially with their perspective, as seen through the weak powers of a good compound Microscope that I am going to order one”.17

16  Letter of October / December 1846 from Darwin to William Benjamin Carpenter, in: Darwin: Correspondence, Vol. III (1987), no. 1001, p. 344. 17  Letter of December 1846 from Darwin to Joseph Hooker, in: Darwin: Correspondence, Vol. III (1987), no. 1035, pp. 375–76, here 375.

VI.  cor al: Tr adition and Encounter

1. The art of transformation Darwin’s favouring of coral as a model for evolution as he had come to understand this would seem to be explained, above all, by the capacity of this organism to “metamorphose” into the petrified stems and arms of its fossil form, its dead and its living manifestations co-existing in a totality composed of two fundamentally distinct entities. But there may well have been further reasons for this choice, relating to the privileged place of coral in the tradition of Natural History. From European Antiquity onwards particular significance had been imputed to coral because it was understood to embody the natural world in its entirety, in addition to that world’s capacity for astonishing feats of transformation. When submerged in water, coral was encountered as a soft and shimmering substance; but, on leaving that element, it hardened into a precious stone. Equally at home in water and in air, coral seemed truly to belong to both. In the second decade AD, in his encyclopaedic compendium Naturalis Historia [On Natural History], the Roman philosopher Pliny the Elder made several references to this dual nature of coral.1 Around seventy years earlier the Roman poet Ovid had marvelled at coral’s capacity for transformation in the oft-quoted lines from Book IV of his Metamorphoses: “And even till this day the same quality has remained in corals, so that they harden when exposed to air, and what was a plant twig beneath the sea is turned to stone above”.2

1 

Pliny, Natural History (ed. 1938 et seq.), Vol. IV. Lib. XII.li, 142, pp. 182 / 183. Vol. VIII, Lib. XXXII, xi, 21–24, esp. 22, pp. 476 / 477–478 / 479. Vol. X, Lib. XXXVII, lvi, 153, pp. 288 / 289. 2  Ovid, Metamorphoses (ed. 1977), Vol. I, Book IV, l. 750–752, pp. 230 / 231: “nunc quoque curaliis eadem natura remansit, / duritiam tacto capiant ut ab aere quodque vimen in aequore erat, / fiat super aequora saxum”. Fundamental on the iconography of coral: Frontisi-Ducroux 1996. A superbly documented account of the history of coral up to the eighteenth century is to be found in Schouppé (1991).

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VI. coral: Tradition and Encounter

37 _ Frontispiece to the volume Museum Richterianum, with commentary by Johann Ernst Hebenstreit, published in Leipzig in 1743

Because coral was thus revealed as itself an artist in metamorphosis, it encouraged primordial forms of awareness of change in the natural world that can, in retrospect, be seen as precursors of the insights that would be achieved in the nineteenth century. Coral was also among the entities most sought after by students of mineralogy. At the same time, it was especially favoured by artists as the raw material from which they might concoct their own extraordinary feats of transformation. When elaborated by those with sufficient skill, it belonged among the most valued of artistic treasures. On occasion, it would be exquisitely combined with other remarkable and precious naturally occurring entities, such as nautilus shells.3 Anyone who has had a chance to visit pre-modern collections embracing both remarkable natural forms and works of art will certainly have come across examples of coral. As revealed by the frontispiece to a mid-eighteenth-century volume on the celebrated Leipzig Museum Richterianum (Fig. 37), coral was among the most

3 

Syndram 1999, p. 133.

1. The art of transformation

valued exhibits in such collections. Prominently positioned in this enticing image, it stood as a pars pro toto, a token of the entire assemblage and, as such, a guarantee of its value.4 Coral signified the profusion of nature in its own capacity as an artist, bountifully offering itself to mankind in its petrified form. And, like the rainbow, coral was a symbol of nature finally at peace with humanity. It is for this reason that

38 _ Francesco Morandini, Allegory of Water. Detail from frescoed ceiling of studiolo of Francesco I de’Medici, 1571. Florence, Palazzo Vecchio

it is to be found spilling forth from a horn-of-plenty in a late sixteenth-century allegory of the Element of Water: Francesco Morandini’s ceiling fresco for the studiolo of Francesco I de’ Medici in the Palazzo Vecchio in Florence (Fig. 38).5 Allusions to coral as a symbol of reconciliation are also to be found in the political iconography of that period. Benvenuto Cellini’s Perseus, erected in 1558 in the Florentine Piazza della Signoria, defiantly holds aloft the severed head of the Medusa (Fig. 39); and the blood oozing from the neck of this hard-won trophy is shown as if suddenly frozen. The artist’s contemporaries would have recognised in this detail an echo of a particular narrative motif from Classical Mythology, which offered a playful explanation for the origin of coral. Perseus, it was claimed, had at one point laid the severed head down upon some seaweed; and this, in reaction to the petrifying power of the Medusa’s gaze, had thereupon hardened. The sea 4 

Bredekamp 2000, pp. 83–84. Dezzi Bardeschi 1980, p. 44. On the allegorical programme of the decoration of the studiolo, see Feinberg 2002. 5 

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VI. coral: Tradition and Encounter

39 _ Benvenuto Cellini, Perseus with the severed head of the Medusa (detail). Bronze. 1558. Florence. Loggia dei Lanzi

nymphs, fascinated and delighted at such a transformation, had then tried this trick for themselves, and had at length scattered the miraculously petrified plant throughout the world’s oceans, where it was to remain: a precious stone for the delight of all. Cellini, drawing together the motif of the trophy set down upon the seaweed, the petrification of this last, the delighted response of the sea nymphs, and the role of coral as a symbol of nature’s bounty, gives the sculpted form of the blood oozing from the Medusa’s severed head the appearance of coral: a token of the former war between mankind and nature now transformed into peace sealed with the presentation of a precious gift. And the bronze Perseus, whose martial stance still threatens all-conquering death to every enemy, thus now also bears witness to a pacific “Age of Coral” in the Florence of the Medici.6 On account of its capacity to fare equally well in water and on dry land, coral offers a counter-image to the destructive forces of nature. In the aforementioned words of Ariel’s song to Ferdinand in Shakespeare’s Tempest (first performed, in London, in 1611) the transformative power of the sea, embodied in coral, is associated with the overcoming of death. For here, the bones of one whom Ferdinand fears to have been drowned in the eponymous storm (his father, King Alonso) are fancifully invoked as already turned to coral –7 although, as the play’s audience is by this point aware, the king and his entourage have in fact been saved from drowning. In Pietro da Cortona’s almost exactly contemporary painting The Coral Fishers 6  7 

Cole 1999, pp. 228–30. See epigraph on p. ix.

2. The cult of coral and the vogue for aquaria

40 _ Pietro da Cortona, The Coral Fishers, c. 1619–21. Catherine Palace, Tsarskoye Selo

(Fig. 40) one finds nothing of this sombre tone, as perhaps befits the character of its first owner, the scholarly Roman naturalist and iconographer Cassiano dal Pozzo (from whose celebrated “paper museum”, the Museo Cartaceo, exquisite images of coral were by no means absent). The emphasis here is on a more elemental form of reconciliation: a triton is seen handing a piece of coral up to a wood nymph as a symbol of the marriage of the Sea and the Earth. This coralline gift, deriving from the subaqueous realm of nature, attests to its true quality in that, upon emerging into the realm of air, it does not expire, but assumes a new and enduring splendour.8

2. The cult of coral and the vogue for aquaria During the mid-nineteenth century the cult of coral underwent diverse forms of intensification. One of the most popular publications of the era was Carl Gottfried Wilhelm Vollmer’s four-volume series of 1854–61, Der Erdball und seine Naturwunder [The Globe and its Natural Wonders]. Illustrated with numerous woodengravings, this went through over fifty editions and was translated into many languages. The third volume, Die Wunder der Urwelt [The Wonders of the Primeval World], initially published in Berlin in 1855, had a section on coral.9 Readers embarking on this account of an ostensibly frail natural form residing in the ocean 8  9 

Solinas 2001, pp. 237–39. Vollmer (1855), 1881, pp. 211–30 , esp. 211–12, 221–25.

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were soon swept up by the author’s veritable hymn of praise to the tirelessly labouring polyps, heretofore largely regarded as a lower form of life. There was here, however, an implicit political message, for scientists had yet to resolve whether each coral formation should be regarded as an assembly of numerous individual entities or as a truly “corporate” organism. This updated version of the political iconography associated with coral is also implicit in Jules Michelet’s volume of 1860 La Mer [The Sea], which decisively contributed to a re-visioning of mankind’s age-old fear of the ocean as a form of worship for the embodiment of Mother Nature at her most tender and all-embracing. In the fourth part of the second and longest section of this text, “La Genèse de la Mer”, Michelet came close to introducing a new cult of coral. For here he never tires of finding in this natural treasure a new and fitting object for his own faible for shadowy natural forms, not least on account of coral’s insistently communal identity. Coral, for Michelet, offers an even more intense promise of happiness because it partakes of all three realms of nature: “There are some ambiguous creatures, the corallinae for example, to which all three realms of the natural world lay claim. They have something about them of the animal; and yet also something of the mineral; and they have of late been deemed to be even more truly vegetal. And perhaps they might be said to occupy that sombre intermediary state, where life arises from its long stony slumber without yet freeing itself entirely from this rugged origin”.10 In the petrified stem of a coral formation Michelet sees a natural strength not deriving from the antagonism of species versus species, but evoked by the virtues of commonality. Mankind’s encounters with coral, it seems to him, serve “to alert us – we who are so proud and who occupy so elevated a perch in the terrestrial fraternity – to the right of the humble mineral world to arise and take on life, and of the deep hopes that dwell within the breast of Nature”.11 To Michelet’s inspired vision, coral is no less strikingly remarkable on account of its role as an indefatigable labourer, capable of erecting gigantic works of architecture through a communal effort – truly noble forms that are suited to its own society: “Within this phantasmagoria, with a little more gravity, the unfurling coral reef displays its somewhat faded colours. Its beauty lies in its form, in this

10  “Il est des êtres incertains. les corallines, par exemple, que les trois règnes se disputent. Elles tiennent de l’animal, elles tiennent du minéral; finalement elles viennent d’être adjugées aux végétaux. Peut-être est-ce le point réel où la vie obscurément se soulève du sommeil de pierre, sans se détacher encore de ce rude point de depart […]” (Michelet 1861, p. 139). 11  “[…] comme pour nous avertir, nous si fiers et placés si haut, de la fraternité ternaire, du droit que l’humble minéral a de monter et s’animer, et de l’aspiration profonde qu’est au sein de la Nature” (Michelet 1861, p. 139).

2. The cult of coral and the vogue for aquaria

form in its entirety, in the noble aspect of the shared city. Each of its inhabitants is modest, but their republic is imposing.”12 The architectural forms of the coral reefs are so very extraordinary because, unlike the cells of bees or humanity’s own works of art, they are not physically external to their creators; rather, the “stones” used to create such marvels are the petrified bodies of the coralline artists themselves, more comparable in this respect with the petrified remains of Herculaneum or Pompeii, albeit without so grim a provenance. The transformation of the bodies of the workers into their own works of art occurs “without violence and without catastrophe, through a natural form of progress; there is a serene peace, an extraordinarily appealing gentleness. Every sculptor would admire the forms achieved here by so marvellous an art”.13 At around this time the cult of coral also played its part in the emergence of a true passion for aquaria. These had, in all probability, first been introduced in Germany in the mid-eighteenth century. And it was there, as also in France, England and Scotland, that efforts were made to ensure that the water they contained was clean and the organisms collected therein healthy. By the mid-nineteenth century, aquaria had become a standard feature in public zoological displays.14 In the eyes of astonished spectators, who had become used to regarding an aquarium as a context for the objective analysis of living organisms, there now suddenly emerged – with the adjunct of coral – a world-in-miniature steeped in bizarrely beautiful colours, which in its secluded self-sufficiency seemed to obey a sort of narcissistic aesthetic. This was, in turn, to become one of the models for the art of the years around 1900.15 Gustave Moreau’s image of Galathea, painted in around 1880 (Fig. 41), constitutes a high-point in the transfiguration of a realm that is in part a gigantic aquarium (such as had indeed been commissioned for inclusion in the Paris Exposition Universelle of 1867) and in part a living coral.16 The nereid is enthroned as if she were being viewed in a glass tank, into which the figure of Polyphemus peers at the upper left, through the third eye located in his forehead. Galathea’s subaqueous throne is decorated with coral formations, and on her head she wears a coral crown entwined with ivy. It is as if Moreau’s image were visually attesting to Michelet’s

12  “Dans cette fantasmagorie, avec plus de gravité, le madrépore arborescant montre ses couleurs moins vives. Sa beauté est dans la forme. Elle est dans l’ensemble surtout, dans le noble aspect de la cité commune; l’individu est modeste, et la république imposante” (Michelet 1861, p. 142). 13  “[…] tout s’est fait sans violence et sans catastrophe, par un progrès naturel; il y a une paix sereine, un astrait singulier de douceur. / Tout sculpteur y admirait les formes d’un art merveilleux” (Michelet 1861, p. 143). 14  Rehbock 1980, pp. 523–34. 15  Harter 2002, Le Paradis, pp. 92–94; Harter 2004. 16  Gustave Moreau 1998, pp. 169–72 , cat. no. 82.

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41 _ Gustave Moreau, Galathea. Oil on board, c. 1880. Paris, Musée d’Orsay

3. Darwin’s own image of coral as an artist

recommendation that the allegorical rendering of Nature always take a form acknowledging that “element of the magical inherent to her” and, in particular, his proposal that Nature be envisaged as enthroned in coral: “Her first-born, the coral reefs, […] would then serve as her seat, extending for this purpose their sprigs, their meanders, their stars of alabaster”.17

3. Darwin’s own image of coral as an artist The diagram that Darwin included in the first, 1859 edition of the Origin of Species fits seamlessly into this older and broader tradition in the appraisal of coral. And it would be hard to underestimate the fact that he had affirmed his appreciation for this larger context not only, by implication, in his own sketches and drawings, but also, both overtly and compellingly, in some of his own earliest published writings. This was already the case in his second publication after returning to England from the H.M.S. Beagle expedition: his monograph of 1842, The Structure and Distribution of Coral Reefs (which appeared as the first part of the “Geology of the Voyage of the Beagle”). Here, Darwin gave voice to his growing conviction that the driving force of nature was expressed in the struggle for life and adaptation, issuing now in defeat, now in victory. But here, and for the first time, he observed that this very struggle also gave rise to natural forms of overpowering beauty.18 The featured wood-engravings of two sites in the south-western Pacific Ocean – Whitsunday Island (Fig. 42, now Pinaki in the Tuamotu Archipelago) and Bolabola (Fig. 43, now Bora-Bora in the Society Islands) – come nowhere near, as Darwin himself conceded, to conveying the exhilaration of which he writes: “[…] every one must be struck with astonishment, when he first beholds one of these vast rings of coralrock, often many leagues in diameter, here and there surmounted by a low verdant island with dazzling white shores, bathed on the outside by the foaming breakers of the ocean, and on the inside surrounding a calm expanse of water, which, from reflection, is of a bright but pale green colour”.19 Darwin was, however, even more impressed by the miraculous allure of the soft, live corals: “The naturalist will feel this astonishment more deeply after having examined the soft and almost gelatinous bodies of these apparently insignificant creatures, and when he knows that the solid reef increases only on the outer edge, which day and night is lashed by the breakers of an ocean never at rest. Well did François Pyrard de Laval, in the year 1605, exclaim “C’est une merueille de voir

17 

“[…] la féerie triomphale qui ne la quitte jamais […] Ses premiers-nés, les madrépores, […] heureux de s’enterrer dessous, en auraient fourni les assises, y mettant leurs rameaux d’albâtre, leurs méandres et leurs étoiles” (Michelet 1861, p. 144). 18  This aspect of Darwin’s writings is developed by Hyman 1974, p. 22. 19  Darwin: Coral Reefs (1842), p. 1.

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42 _ Whitsunday Island (now Pinaki, Tuamotu Archipelago) in the south-western Pacific Ocean. Illustration (from an earlier publication) in Darwin’s volume of 1842 The Structure and Distribution of Coral Reefs

chacun de ces atollons, envionné d’un grand banc de pierre tout autour, n’ayant point d’artifice humain”.20 The reference to Pyrard de Laval reveals that Darwin saw himself not only as the destroyer of an intellectual tradition, but also as an eager contributor to its by no means exhausted discourse. This becomes even clearer in the admiration expressed in his diary (compiled at intervals, while still at sea, as a continuous narrative derived from daily notebook entries) for the Prussian geologist, botanist and intrepid explorer Alexander von Humboldt, and in particular for Humboldt’s ability to convey a sense of his own informed exhilaration at the natural world. The entry for 28th February 1832, written shortly after the Beagle reached Bahia, on the coast of Brazil (where Darwin was at last able to immerse himself in lush surroundings of the sort that recalled Humboldt’s account of his own Central and South American travels of 1799–1804) contains the acclamation: “Humboldts glorious descriptions are & will for ever be unparalleled […] the rare union of poetry with science”;21 and the confession that Humboldt was, for him, “like another Sun”.22 And, over a decade

20 

Darwin: Coral Reefs (1842), pp. 1–2. Cited French text reads: “It is a wonder to behold each of these atolls, surrounded as it is by a great bank of stone created entirely without human artifice”. The travels of François Pyrard de Laval (c. 1570–1621), taking in Brazil, the Indian Ocean and the East Indies, spanned a decade, 1601–11. A first account appeared in Paris in 1611, with several further editions following. 21  Darwin: Beagle Diary (ed. 1988), pp. 41–44, here 41. Cf. Leask 2003, p. 23. The 1988 edition is an exact transcript of the 751-page manuscript covering the events of 24 October 1831 to 7 November 1836. 22  Darwin: Beagle Diary (ed. 1988), p. 42. Cf. Leask 2003, p. 25). This may also be seen as a reason for Darwin’s receptivity to metaphor, an issue superbly developed in: Beer 1983, pp. 89–98, 103.

3. Darwin’s own image of coral as an artist

43 _ Bolabola (now Bora-Bora, Society Islands) in the south-western Pacific Ocean. Illustration (from an earlier publication) in Darwin’s volume of 1842 The Structure and Distribution of Coral Reefs

later, in 1845, Darwin was still to be found recalling that he had, as a young man, read Humboldt’s “Personal Narrative” (title of the first English translation of the account of his travels) and that this had had a huge impact on the decisive turn that his own life had then taken.23 Darwin’s endeavour to blend a conventional English restraint with the style of Humboldt, as rendered by his translator, was perhaps at its most successful in the record of his only direct encounter with coral reefs, in the south-western Pacific, and in particular the Indian, Oceans during the last year of the circumnavigatory voyage. This is to be found in the penultimate chapter of his first related publication, of 1839, the Journal and Remarks, 1832–1836 (the third volume of the official

23  “I never forget that my whole course of life is due to having read & reread as a Youth his Personal Narrative”. Letter of 10 February 1845 from Darwin to Joseph Hooker, in: Darwin: Correspondence, Vol. III (1987), no. 826, pp. 139–40, here 140. Cf. Leask 2003, p. 16, and passim., on Darwin’s life-long reverence for Humboldt. The latter’s Central and South American travels, in the company of his French colleague Aimé Bonpland, between 1799 and 1804, armed him with the extensive experience and data on which he drew during his subsequent long sojourn in Paris (1804– 27) for the thirty volumes initially published there, in French, between 1805/07 and 1834. Of these, the more widely appealing three-volume Relation historique du Voyage aux régions équinoxiales du Nouveau Continent […], published in 1814, 1819 and 1825, (see Humboldt 1814–25), was translated by Helen Maria Williams, a largely expatriate Englishwoman with a pre-established literary career and her own strong interest in Central and South America, who embarked on her collaboration with Humboldt in around 1810. Her somewhat flowery and over-dramatised translation, in seven volumes, was published in London between 1814 and 1829; see Humboldt (1814– 29).This is the “Personal Narrative” to which Darwin referred. A copy of the third edition of volume one had been presented to him, by John Henslow, in September 1831, shortly before H.M.S. Beagle set off on its circumnavigatory voyage, affectionately inscribed in acknowledgement of the significance this new episode would hold for him.

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“Narrative of the Surveying Voyages of […] 1826–1836”).24 It largely reflects upon what he was himself able to observe and examine in the chief atoll of the Keeling (Cocos) Islands (Fig. 44).25 Of most interest to him here was the ring of coral rock, above all its ocean-facing, live-coral exterior. Struck by the terrifying force of the ocean perpetually breaking against the reef, Darwin is initially prepared to see this as “an invincible, all-powerful enemy”. But, upon reflection, and in a spirit that anti­­cipates both Vollmer and Michelet, he finds the more remarkable natural phenomenon to be the ultimate resistance to the destructive “hurricane” achieved through the tireless activity of the coral-building polyps: “what will this [destructive force] tell against the accumulated labour of myriads of architects at work day and night […] Thus do we see the soft and gelatinous body of a polypus, through the agency of the vital laws, conquering the great mechanical power of the waves of an ocean, which neither the art of man, nor the inanimate works of nature could successfully resist”.26 As a result of this assiduous activity, performed in countless, infinitely slow, infinitely tiny, yet at length extremely effective steps, coral succeeds in erecting vast constructions that, in Darwin’s view, are far more imposing than the greatest ruins from Antiquity.27 Darwin finds further evidence of coral illustrating the capacity of “nature as an artist” in his reflections on some of the more complex coral formations. He marvels in particular at the sight of an “encircling reef”, in which the sheltered coral lagoon has an island at its centre: “Can any thing be more singular than this structure? It is analogous to that of a lagoon, but with the island standing, like a picture in its frame, in the middle”.28 If, in Darwin’s view, coral and other natural forms brought forth through the processes involved in the struggle for life,

24  ”Narrative of the Surveying Voyages of His Majesty’s Ships Adventure and Beagle, between the Years 1826–1836, describing their Examination of the Southern Shores of South America, and the Beagle’s Circumnavigation of the Globe”, London 1839. 25  As revealed by Darwin’s comments on the ten maps illustrated on the fold-out sheet prefacing his volume on the Coral Reefs (most of them derived, as he explains, from those produced, in several “very different styles”, following diverse earlier surveying expeditions), his observations on that of the Keeling (Cocos) Islands (p. ix) are unusually vivid and detailed: evidence of his personal experience of the site, enriched by the expertise of his accomplice there, Robert FitzRoy, to whom this particular map is owed. 26  Darwin: Journal and Remarks (1839), p. 338. 27  Darwin: Journal and Remarks (1839), p.553: “I am glad we have visited these islands: such formations surely rank high amongst the wonderful objects of the world. It is not a wonder, which at first strikes the eye of the body, but rather, after reflection, the eye of reason. We feel surprised, when travellers relate accounts of the vast extent of certain ancient ruins; but how utterly insignificant are the greatest of these, when compared to the pile of stone here accumulated by the work of various minute animals.” 28  Darwin: Journal and Remarks (1839), p. 555.

3. Darwin’s own image of coral as an artist

44 _ The Keeling (Cocos) Islands in the Indian Ocean. Map included (as part of Plate I) in Darwin’s volume of 1842 The Structure and Distribution of Coral Reefs

belonged in the realm of the work of art, then he can be seen as operating within an intellectual tradition that was still alive well into the nineteenth century.29 Of certainly no less significance for Darwin was the role of coral as an embodiment of “pacification” in the conflict between the power of the sea and that of the land. In retrospect, after his return to England, as he pondered what he had himself observed and understood of the coral reefs, these elongated edifices built up over thousands of years, and the outcome of countless minute organic processes, may well have come to resemble “monuments” erected in acknowledgement of the ultimate irresistibility, yet infinite slowness, of evolution in the natural world: the contribution of each living entity in the coral polypifer being infinitely tiny and yet, cumulatively and over a vast span of time, superior to any form of mechanical power.

29 

Stafford 1976.

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For all the scientific sophistication he was eventually able to bring to the phenomenon of coral, Darwin never ceased to see in it the qualities with which it had been associated in pre-modern Natural History. For him it remained a symbol of all that was alive, a symbol arising out of the conflict between sea and land, and which, through its own activity, as a tireless “artist”, was capable of preparing an earthly paradise for mankind. Darwin was eventually to confess that he felt his own effective destruction of the biblical account of Creation had been tantamount to “a murder”.30 But, to the struggle between the species for their own survival – a struggle (he conceded) no less terrible than it was (as he also insisted) exalting – coral seemed to offer, as it had for centuries, indeed millennia, a true alternative. It was in the summer of 1858 that Darwin embarked on the shorter version of his manuscript on “natural selection” that was to become the published text of his Origin of Species. At almost exactly the same time Michelet was composing his hymn to the sea and to coral. In some respects the two may well strike one as virtual opposites: while Michelet still evoked the world as it had long been understood in terms of traditional Natural History, this perspective was in due course to retreat, owing not least to the scientific contribution of Darwin. Ultimately, however, such a view of these close contemporaries is too simplistic. For, thanks to the intensity of his feeling for coral, nature as perceived by Darwin remained an aesthetically enchanted world.31

30  Letter of 11 January 1844 from Darwin to Joseph Hooker, in: Darwin: Correspondence, Vol. III (1987), no. 729, pp. 1–3, here 2: “I am almost convinced (quite contrary to [the] opinion I started with) that species are not (it is like confessing a murder) immutable”. Cf. Hyman 1974, p. 30. 31  In as far as Evolutionary Biology has “forgotten” this aspect of Darwin’s relationship to nature, it may effectively be seen to have incurred that intellectual impoverishment in which the contest between Evolutionists and Creationists persists.

Conclusion: The lure of the “endless” Darwin’s awareness of coral was at first gradual and haphazardly accumulative. But it was soon to attain a far greater sophistication as chance discoveries encouraged new perspectives, and these in turn revealed to him the deeper significance of knowledge imbibed earlier. Having all but effortlessly absorbed the age-old cultural and epistemic associations of coral (both as a token of “metamorphosis” in the natural world, and as an epitome of nature in its entirety), Darwin was to bring these to bear on the often searingly vivid experiences that came his way on the Beagle expedition, several of which featured coral or related forms. During those five years he was to be found no less keenly engaged, on the bleak shores of Patagonia, enquiring into all that might be gleaned from the most intense scrutiny of a single specimen (of the sort reflected in the later notes on his find of January 1834, Fig. 1)1 than he was eloquently ecstatic in evoking his response to the long-awaited opportunity to examine a coral reef (in the diary entries for November 1835 and, in particular, April 1836, on which he drew for his publications of 1839 and 1842).2 With a now far greater understanding of coral in all its manifestations, Darwin was able fully to appreciate its astounding “constructive” capacity and to grasp both the infinite delicacy of each organic contribution to the entire operation and the vast chronological scale over which this occurred. He was also able to perceive that there was a heartening symbolism to be found in the emergence of such ravishing natural beauty out of a ferocious and relentless battle between land and ocean. These insights, as they recurred to Darwin after his return to England, were in due course to be corrected, adapted, enhanced or elaborated through all that he was able to learn or surmise by reference to an extensive range of older and newer literature, an assiduously cultivated network of accommodating scholarly correspondents, 1 

See Darwin: “Coralline Algae Notes” (ed. 1987), p. 194. See, in particular, Darwin’s vivid account of his experiences of 6 and 12 April 1836, during his aforementioned time in the Keeling (Cocos) Islands, in: Darwin: Beagle Diary (ed. 1988), pp. 417–18. 2 

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and a number of tirelessly helpful colleagues and friends. In the light of so much new input, it is telling that, in his thinking on species transmutation, the three distinct sorts of visual model to be committed to paper and preserved – the coralline sketches of 1837 (Fig. 6, 14), the circle segment drawings of the early and mid1850s (Figs. 20, 23, 24), the diagrams of 1857–59 (Figs. 29, 33) – all retain features that derive from notions first sparked, if not yet seriously entertained, during his most fruitfully “impressionable” years. Prominent among these are: a firm sense of an entity functioning as an emphatically bi-partite whole, some of it living, most of it dead (Figs. 6, 33); development in the form of “sprawling”, multi-directional ramification, with a strict balance of “success” and “failure” (Figs. 14, 33); the potential co-existence of undivergent and highly divergent strains (Fig. 20) or of radically contrasting patterns of mutation (Fig. 23); adjustment to a time-frame of mindstretching immensity, entailing an “interminable” present suspended between virtual “infinities” of past and future (Fig. 33). * However well justified our focus upon coral, it is nonetheless salutary to consider this also in the light of another, not entirely unrelated natural form to which Darwin looked in pursuit of alternatives to the tree as the inspiration for a compelling model to epitomise his theory. At the top of the verso of one of around three hundred loose sheets (some torn from notebooks) gathered by Darwin in a portfolio of material relevant to his thoughts on species transmutation, is the impetuously pencilled assertion “a tree not good simile”, directly followed by a dash, then the naming of the here favoured substitute: “endless piece of sea weed dividing”. This statement is, moreover, accompanied by a cursory pencil rendering of what Darwin has in mind (Fig. 45): an image remarkable not only on account of its visual echo of his Patagonian specimen (Fig. 1), but equally as proof that the abstract essence of such an organism was sufficiently firmly lodged in Darwin’s mind to afford even this weak draughtsman effortless manual access to it. The comment on the verso of the sheet is also evidently a reactive continuation of a line, also in pencil, that Darwin has appended, as an explanatory aside, to the opening of his principal text, written in ink, on the recto, which is dated, at the upper left and in his own hand (though possibly in retrospect), “July 1843”. That text, though lacking in grammatical cohesion, is clear in its thrust: Darwin here observes that, just as “all groups in my theory blend with each other”, the unidirectional relation of a larger to a smaller category (a family to a genus, a genus to a species, a species to a variety) cannot be assumed to remain stable indefinitely, for to assume so would be to misrepresent nature as it truly is. (As the reader will recall, this was the period of Strickland’s greatest influence on Darwin. And, as Darwin’s explication of the 1859 diagram was to affirm, varieties, when sufficiently distinct, would themselves qualify as new species.) In illustration of this “blending”, Darwin

Conclusion: The lure of the “endless”

45 _ Sketch accompanying a note on the suitability of seaweed as a model for evolution (species transmutation) through a process of “natural selection”. Pencil on paper, 1843. Cambridge University Library, DAR 205.5: portfolio of loose sheets, fol.  90v. Reproduced with the permission of the Syndics of Cambridge University Library and of William Huxley Darwin

in his explanatory aside embarks on citing the concrete example of a tree, in which “no part or branch […] can be said to be distinct”. Such a “simile” then striking him as inappropriate for this purpose, the “endless piece of sea weed” invoked on the verso would here have appealed for the very ease with which imagination can find in it not only an emblem of the disorderly profusion of nature, but even an intimation of indivisibility and infinity.3 * Those familiar with the rousing conclusion to the Origin of Species may have guessed that the term “endless” was a particular favourite of Darwin’s and, moreover, that he invested it with far more positive connotations than would be usual for a native speaker of English in the mid-nineteenth century, as indeed in the early twentyfirst. It would thus be irresponsible to over-state the secondary meaning of the term “end”, signifying “goal” or “purpose”: for then (as now) the use of “endless” to signify “with no end in sight” was by no means equivalent to the use of “end-less” to 3  Darwin indicates that his exposition continues from recto on to verso through his habitual use, at the lower right of the former, of a short diagonal stroke placed above the term “over”. Gruber 1988, p. 127 recalled noting the inscription regarding seaweed among material viewed at Cambridge University Library, but failed to supply a specific location for this. He did not identify the drawn form as intended to represent seaweed, having already stated his assumption that all Darwin’s sketches of this sort showed “trees”. Nor did he observe the explicit interconnection of the text on the recto with that on the verso of the sheet.

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signify “without purpose; pointless”. But it is clear (not least if one recalls the nature of Darwin’s earlier approval for the “endless piece of sea weed”) that more than a hint of ambiguity persists within his closing paean to nature’s “endless forms most beautiful and most wonderful”.4 As of 1859 an essential “pointlessness” lay at the heart of Darwin’s vision of the natural world, of how its forms had evolved, and of how they would go on and on and on evolving. And it is incumbent upon every blithely retrospective engagement with his work to be vigilant in the effort to recall precisely how dispiriting Darwin’s theory could seem to many, indeed most, in mid-nineteenth-century England. It is, however, no less imperative to acknowledge that Darwin also attended to what he saw in, or rather through, the “pointless” struggle for life and for survival: a countervailing impetus, which drew forth beauty from the superfluity of energies released in that struggle. This was a phenomenon he had very consciously beheld at its most stirring among the coral reefs of the Pacific and Indian Oceans: an experience he readily identified as the highpoint of a life-changing and life-defining episode. It is above all Darwin’s eye for the beauty to be discovered in nature (a “purpose” for which was, in his later work, to eclipse the spectre of “pointlessness”)5 that reveals him as a Janus-headed genius: both the destroyer and the completer of a tradition of Natural History. While Darwin was at length to become a popular “classic” among nineteenth-century thinkers and writers, in his perceived role as an out-and-out revolutionary, it is ultimately in the preservation of tradition that one may find his more truly superlative achievement.

4 

Darwin: Origin of Species (1859), p. 490. This was very much the case in Darwin’s volume of 1871, The Descent of Man and Selection in Relation to Sex, in wich “sexual selection” – ultimately on the basis of the perceived “beautiful form” of a preferred mate – is proposed as an evolutionary mechanism complementary to that of “natural selection” as expounded in the Origin of Species. 5 

Acknowledgements For the always generous provision of assistance – be it in the form of information, advice, or graciously supplied correction – I am much indebted to Carmen Alonso Schmitt, Mario A. Di Gregorio, Ulrike Feist, Stephen Greenblatt, Michael Hagner, Ursula Harter, Nadine Helm, Wolfram Hogrebe, Ulf Jensen, Rebekka Maiwald, Amy Meyers, Gerhard Neuweiler, Magdalena Popp-Grilli, Henning Ritter, Helmut Rohlfings, Martin Rudwick, Gerhardt Scholtz, Steffen Siegel, Esther Sturm, Rüdiger Wehner, and Reinhard Wendler. My particular thanks are due to Barbara Herrenkind for her outstanding photographic skills, and to the determined and resourceful librarians of the Wissenschaftskolleg zu Berlin, who tracked down many an obscure publication. I am especially grateful to Adam J. Perkins, Curator of Scientific Manuscripts, Cambridge University Library, to Jennifer Bryant and Linda M. Irvine of the Natural History Museum in London, and to Carsten Lüter of the Museum für Naturkunde in Berlin for the interest and enthusiasm they evinced in granting me access to the precious material in the archives and collections in their care. And it is to Till Kreische that I owe the happy suggestion of the lines from Shakespeare that here serve as my epigraph. My greatest debt is, however, to Julia Voss. While I was deeply absorbed in Darwin’s coral iconology, she was bringing to completion a comprehensive study of the visual character of Darwin’s thought and its repercussions: an ambitious project that was to issue in the first analysis of Darwin’s visual world to draw equally on Art History and the History of Science.1 It is from this scholar – through her encouragement no less than through her assistance – that my own work on Darwin has profited the most. The editorial staff of Wagenbach Verlag in Berlin, publisher in 2005 of this book its original, German version, Darwins Korallen. Die frühen Evolutionsdiagramme

1  Voss 2005. This doctoral dissertation was subsequently published in its original German version (see Voss 2007), and then in English translation (see Voss 2010).

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und die Tradition der Naturgeschichte, brought their characteristic sensitivity and flair to the presentation of my theme. As so often in the past, I am pleased to extend my great thanks to Petra Florath, whose talents are here much in evidence in the reconciliation of the textual and illustrational aspects of the book so as to accommodate an unusually wide range of images. My collaboration with Elizabeth Clegg on the English adaptation of my text has proved fruitful even beyond the expectations raised by our earlier projects. No less valued than her gift for detecting, then deftly repairing, the more serious lacunae in my original exposition has been her guidance through the numerous important additions to the field of Darwin studies since 2005, both publications and online resources.2 All the more have I been heartened by her assurance that my own contribution still has much to offer the far wider readership to which it can now be addressed.

2  For the greater convenience of readers, the post-2005 bibliography compiled as an adjunct to the adaptation has been merged with an extended version of the original, 2005 bibliography in a single, alphabetically ordered list.

Bibliogr aphy Where several publications by the same author are cited, these appear in chronological order of their publication. This bibliography is restricted to the original-language version of each item cited, and has not (with certain exceptions) been further extended through the addition of existing English translations of non-anglophone texts. Abeles, Oren, “The agricultural figures of Darwin’s evolutionary rhetoric”, The Quarterly Journal of Speech, CII / 1 (2016), pp. 41–61 Agassiz, Louis and A. A. Gould, Outlines of Comparative Physiology […], Boston, Mass. 1848; enlarged edn. Boston and London 1851 Alberti, Leon Battitsta, De Statua, Latin text with English trans. by Mariarosaria Spinetti, Naples 1999 Allbach, Horst and Erwin Nehr, eds., Alexander von Humboldt und Charles Darwin: Zwei Revolutionäre wider Willen, Göttingen 2011 Allmon, Warren D., “Darwin and palaeontology: a re-evaluation of the interpretation of the fossil record”, Historical Biology, XXVIII / 5 (2016), pp. 680– 706 Alter, S. G., “Mandeville’s Ship. Theistic Design and Philosophical History in Charles Darwin’s Vision of Natural History”, Journal of the History of Ideas, LXIX / 3 (2008), ppp. 441–66 Altner, Günther, Charles Darwin und die Dynamik der Schöpfung: Natur, Geschichte, Evolution, Schöpfung, Gütersloh 2003 Amigoni, David and Jeff Wallace, eds., Charles Darwin’s ‘The Origin of Species’. New Interdisciplinary Essays, Manchester and New York 1995 Aquinas, Thomas, Opuscula, ed. Ottaviano Scoto, Venice 1498 Archibald, J. D., “Darwin’s two competing phylogenetic trees: marsupials as ancestors or sister taxa?”, Archives of Natural History, XXXIX (2012), pp. 217–33 Archibald, J. D., Aristotle’s Ladder, Darwin’s Tree. The Evolution of Visual Metaphors for Biological Order, New York 2014 Ascione, Gina Carla, Storia del corallo a Napoli dal XVI al XIX secolo, Naples1991 Ashton, Rosemany, One hot summer: Dickens, Darwin, Disraeli, and the great stink of 1858, New Haven and London 2017 Barry, Martin, “On the Unity of Structure in the Animal Kingdom”, Edinburgh New Philosophical Journal, XXII (1836/37), pp. 116–41, 345–64 Barsanti, Giulio, La scala, la mappa, l’albero. Imagini e classificazioni della natura fra Sei e ­Ottocento, Florence 1992 Barta, Ilsebill, Familienportraits der Habsburger. Dynastische Repräsentation im Zeitalter der ­Aufklärung, Vienna, Cologne and Weimar 2001

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Becquemont, Daniel, Charles Darwin 1837–1838, aux sources d’une découverte, Paris 2009 Beddell, B.G., “Darwin and divergence: The Wallace connection’”, Journal of the History of ­Biology, XXI / 1 (1988), pp. 1–68 Beer, Gillian, Darwin’s Plots. Evolutionary Narrative in Darwin, George Eliot and NineteenthCentury Fiction, London 1983 Beer, Gillian, “Four bodies on the Beagle: touch, sight and writing in a Darwin letter”, in: Judith Still and Michael Worton, eds., Textuality and Sexuality. Reading theories and practices, Manchester 1993, pp. 116–32 Beer, Gillian, Open Fields: Science in Cultural Encounter, Oxford 1996 Bell, G., Selection: the mechanism of evolution, New York 1997 Bell, Michael A. et al., Evolution since Darwin: The first 150 Years, Sunderland, Mass. 2010 Berns, Jörg Jochen, “Baumsprache und Sprachbaum. Baumikonographie als topologischer Komplex zwischen 13. und 17. Jahrhundert”, in: Kilian Heck and Bernhard Jahn, eds., Genealogie als Denkform in Mittelalter und Früher Neuzeit, Tübingen 2000, pp. 155–246 Blancke, Stefan et al., “From Ends to Causes (and Back Again) by Metaphor: The Paradox of Natural Selection”, Sciences and Education, XXIII / 4 (2014), pp. 799–808 Bogen, Steffen and Felix Thürlemann, “Jenseits der Opposition von Text und Bild. Überle­ gungen zu einer Theorie des Diagramms und des Diagrammatischen”, in: Alexander Patschovsky, ed., Die Bildwelt der Diagramme Joachims von Fiore. Zur Medialität religiöspolitischer Programme im Mittelalter, Ostfildern 2003, pp. 1–22 Bonhoff, Ulrike Maria, “Das Diagramm. Kunsthistorische Betrachtung über seine vielfältige Anwendung von der Antike bis zur Neuzeit”. Doctoral dissertation, Westfälische WilhelmsUniversität, Münster 1993 Bonnet, Charles, Contemplation de la Nature, 2 vols., Amsterdam 1764 Bowler, Peter J., The Eclipse of Darwinism: Anti-Darwinian Theories in the Decades around 1900, Baltimore and London 1983 Bowler, Peter J., Theories of human evolution: a century of debate, 1844–1944, Baltimore 1986 Bowler, Peter J., Charles Darwin, Oxford 1990 Bowler, Peter A., Darwin deleted: imagining a world without Darwin, Chicago 2013 Bowles, Samuel and Peter Hammerstein, “Does Market Theory Apply to Biology?”, in: Genetic and cultural evolution of cooperation, Berlin 2003, pp. 153–65 Brackmann, A.C., A delicate arrangement: the strange case of Charles Darwin and Alfred Russel Wallace, New York 1980 Bredekamp, Horst, Antikensehnsucht und Maschinenglauben. Die Geschichte der Kunstkammer und die Zukunft der Kunstgescichte, Berlin 1993; rev. ed. 2000; trans.(from 1993 edn.) Allison Brown, The Lure of Antiquity and the Cult of the Machine: the Kunstkammer and the Evolution of Nature, Art and Technology, Princeton 1995 Bredekamp, Horst, “Baum und Überbaum. Zur Semantik eines Bildes der Evolution des Lebens”, Frankfurter Allgemeine Zeitung, 150 (2 July 2003), p. N3 Bredekamp, Horst, “Darwins Evolutionsdiagramm, oder: Brauchen Bilder Gedanken?”, in: Wolfram Hogrebe and Joachim Bromand, eds., Grenzen und Grenzenüberschreitungen. XIX. Deutscher Kongress für Philosophie, Bonn 23–27. September 2002. Vorträge und ­Kolloquien, Berlin 2004, pp. 863–77 Bredekamp, Horst, “Die wilde Üppigkeit der Natur. Stricklands Karten und Darwins Kreise der Arten”, in: Natascha Adamowskyand Peter Matussek, eds., [Auslassungen]. Leerstellen als Movens der Kulturwissenschaft, Würzburg 2004, pp. 341–53 Bredekamp, Horst, “Darwins Korallen”, in: Anke te Heesen and Petra Lutz, eds., Dingwelten. Das Museum als Erkenntnisort, Dresden 2005, pp. 77–87 Bredekamp, Horst, “Denkende Hände. Überlegungen zur Bildkunst der Naturwissenschaften”, in: M. Lessl and J. Mittelstrass, eds., Von der Wahrnehmung zur Erkenntnis. From Perception

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to Understanding. Symposium der Schering Forschungsgesellschaft zu Ehren von Professor Dr. Dr. h.c. Günter Stock, Februar 2004, Heidelberg 2005, pp. 109–32 Bredekamp, Horst, “Die zeichnende Denkkraft”, in: Jörg Hubner, ed. Einbildungen (Interven­ tionen:14), Zürich 2005, pp. 155–71 Bredekamp, Horst, “Bilder in Evolution und Evolutionstheorie”, in: Evolution und Menschwerdung: Vorträge anlässlich der Jahresversammlung der Leopoldina von 7. bis 9. Oktober 2005 zu Halle / Saale, ed. Harald zur Hausen, Halle 2006, pp. 195–215 Bredekamp, Horst, “Darwins Korallen und das Problem animalischer Schönheit”, in: Bilderwelten. Vom farbigen Abglanz der Natur, ed. Norbert Elsner, Göttingen 2007, pp. 257–80 Bredekamp, Horst, “Das Prinzip der Metamorphosen und die Theorie der Evolution”, in: BerlinBrandenburgische Akademie der Wissenschaften Jahrbuch 2008, Berlin 2009, pp. 209–47 Bredekamp, Horst, “Das Modell als Fetisch und Fessel”, in: Nova Acta Leopoldina, new series, CXIII / 388 (2012), pp. 61–99 Bredekamp, Horst, “Corals versus Trees: Charles Darwin’s Early Sketches of Evolution”, in: Pamela H. Smith, Amy R.W. Meyers and Harold J. Cook, eds., Ways of Making and Knowing: the material culture of empirical knowledge, Ann Arbor 2014, pp. 357–76 Breidbach, Olaf, “Einleitung”, in: C.E. Baer, ed., Über Entwicklungsgeschichte der Thiere. Beobachtung und Reflexion [1829], Hildesheim and Zürich 1999, pp. V–XXIII Bronn, Heinrich Georg, Untersuchungen über die Entwicklungs-Gesetze der organischen Welt während der Bildungs-Zeit unserer Erd-Oberfläche, Stuttgart 1858 Brooks, John Langdon, Just before the Origin: Alfred Russel Wallace’s Theory of Evolution, New York 1984 Brown, William and Andrew C. Fabian, Darwin, Cambridge 2010 Browne, Janet, Charles Darwin: Voyaging, London 1995 Browne, Janet, Charles Darwin: The Power of Place, London 2002 Browne, Janet, Darwin’s Origin of Species: a Biography, London 2007 Browne, Janet, “Making Darwin: Biography and the Changing Representations of Charles ­Darwin”, The Journal of Interdisciplinary History, XL / 3 (2010), pp. 347–74 Browne, Janet, “Wallace and Darwin”, Current Biology, XXIII / 24 (2013), pp. 1071–72 Bucci, Mario, Lo studiolo di Francesco I, Florence 1965 Burkett, Andrew and Josh Lanier, “Chance in Darwinian Evolutionary Theory and British Romanticism”, Literature Compass, XIII / 10 (2016), pp. 663–89 Campbell, John Angus, On the Way to the Origin. Darwin’s Evolutionary Insight and its Rhetorical Transformation, Evanston, Il. 1990 Campbell, John Angus and Stephen C. Meyer, eds., Darwinian design and public education, East Lansing, Mich. 2003 Canguilhem, Georges, Études d’Histoire et de Philosophie des Sciences, Paris 1994 Cantor, Geoffrey N., and Sally Shuttleworth, eds., Science Serialized: Representation of the Sciences in Nineteenth Century Periodicals, Cambridge, Mass. and London 2004 Carroll, Joseph, Literary Darwinism: evolution, human nature, and literature, New York and ­London 2004 Carroll, Joseph, Dan P. McAdams and Edward O. Wilson, eds., Darwin’s Bridge: uniting the humanities and sciences, New York and Oxford 2016 Carroll, S.B., The making of the fittest: DNA and the ultimate forensic record of evolution, New York 2006 Cohen, Claudine, “How nationality influences Opinion: Darwinism and palaeontology in France (1859–1914)”, Studies in the History and Philosophy of Biological and Biomedical Sciences, LXVIII (2018), pp. 8–17 Cole, Michael, “Cellini’s Blood”, Art Bulletin, LXXXI / 2 (June 1999), pp. 215–35

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Coralli talismani sacri e profani, exh. cat., Trapani – Museo Regionale Pepoli, Palermo 1956; Palermo 1956 Cowles, H.M., “On the Origin of Theories: Charles Darwin’s vocabulary of method”, The American Historical Review, CXXII / 4 (2017), pp. 1079–1104 Crick, Nathan, “’A Capital and Novel Argument’. Charles Darwin’s Notebooks and the Productivity of Rhetorical Consciousness”, The Quarterly Journal of Speech, XCI / 4 (November 2005), pp. 337–64 Cronin, Helena, The Ant and the Peacock, Cambridge, Mass. 1991 Cutler, A., The seashell and the mountaintop, New York 2003 Daneu, Antonio, L’Arte Trapanese del Corallo, Palermo 1964 Darwin, Charles 1. online resources A useful guide to these is to be found in Adam M. Goldstein, “Charles Darwin’s Manuscripts and Publications on the World Wide Web”, Evolution and Outreach, II / 1 (January 2009), pp. 122–35 http://www.darwin-online.org.uk Initiated in 2002, and now extensive in its contents, this is useful for the purposes of rapid ­reference, especially if the searcher is already possessed of a secure identification for the item sought. It embraces both publications (notably, first editions of Darwin’s own published writings) and manuscripts (although the image quality of these is sometimes poor, few ­transcriptions are as yet supplied, and there are no editorial annotations). http://www.darwinlibrary.amnh.org Initiated in 2005 at the American Museum of Natural History (in connection with an exhibition staged there) and re-launched in 2009, this comprises the Darwin Manuscript Project and the Library of Evolution. Although as yet much less extensive than the aforementioned site, it is particularly valuable for the quality of its photographic record of the manuscripts, for the authoratitive transcripts supplied, and for its editorial annotations (which, crucially, attend to the palimpsestic character of this resource). http://www.darwinproject.ac.uk This is the online version of the fruits of the ambitious Darwin Correspondence Project, initiated in 1974, which aims to issue in print all of Darwin’s extant correspondence, the first volume of which appeared in 1985 (see below). 2. correspondence to and from Darwin The Correspondence of Charles Darwin, Frederick Burkhardt, Sydney Smith et al., eds., Cambridge 1985 et seq. (Of the projected total of thirty volumes, Vol. XXV, covering 1877, appeared in 2017.) Volumes cited in this book: I (1985): 1821–36; II (1986): 1837–43; III (1987): 1844–46; IV (1988): 1847–50; V (1989): 1851–55; VI (1990): 1856–57; VII (1991): 1858–59 & supplement for 1821–57 Darwin, Francis, The Life and Letters of Charles Darwin, including an autobiographical chapter, 3 vols., London 1887 Darwin, Francis and A. C. Seward, eds., More letters of Charles Darwin, 2 vols., London 1903 [while these earlier editions are now superseded by the Correspondence volumes appearing since 1985, they are, of necessity, cited in the earlier Darwin literature] 3. further manuscript material in transcribed and edited form (in chronological order of the original manuscript) Charles Darwin’s Notebooks from the Voyage of the Beagle, transcribed and intr. Gordon Chancellor and John van Wyhe, Cambridge 2009 [transcript in full, with excellent commentaties, of the fifteen small field notebooks kept by Darwin in 1832–38]

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Richard Darwin Keynes, Charles Darwin’s Beagle Diary, Cambridge 1988 [transcript of the diary complied by Darwin through adaptation from entries in the above-mentioned notebooks] Nora Barlow, Charles Darwin’s Diary of the Voyage of H.M.S. “Beagle”, Cambridge 1933 [a pioneering work, but now superseded by the previous volume] Nora Barlow, Charles Darwin and the Voyage of the Beagle, London 1945 [partial transcripts of Darwin’s original Beagle notebooks, and of related correspondence; published as a supplement to the same editor’s volume of 1933; now superseded, as regards the notebooks, by the above-cited volume of 2009 and, as regards the correspondence, by the volumes appearing since 1985] Darwin’s “Coralline Algae Notes” [made during and after 1831–36], in: Porter, Duncan M., “Darwin’s Notes on Beagle plants”, in: Bulletin of the British Museum (Natural History: Historical Series) XIV / 2 (26 November 1987), pp. 186–217 Charles Darwin’s Notebooks, 1830–1844, Geology, Transmutation of Species, Metaphysical Enquiries, transcribed and ed. Paul H. Barrett, Peter J. Gautrey et al. (British Museum, Natural History), London 1987 Francis Darwin, ed., The Foundations of the Origin of Species. Two essays written in 1842 and 1844 by Charles Darwin, Cambridge 1909 R. C. Stauffer, ed., Charles Darwin’s “Natural Selection”, being the second part of his Big Species Book (1856–58), edited from the manuscript, Cambridge 1975 Nora Barlow, ed., The autobiography of Charles Darwin 1809–1882, with the original omissions restored, London 1958 [edition of the “autobiographical sketch” that Darwin compiled ­sporadically between 1876 and 1881] 4. Darwin’s own publications mentioned in this book Journal and Remarks. 1832–1836, London 1839 (= Vol. III of “Narrative of the Surveying ­Voyages of His Majesty’s Ships Adventure and Beagle, between the Years 1826 and 1836, describing their Examination of the Southern Shores of South America and the Beagle’s Circumnavigation of the Globe”) The Structure and Distribution of Coral Reefs, London 1842 (= Vol. I of “The Geology of the ­Voyage of the Beagle, under the Command of Capt. Robert FitzRoy, R.N. during the Years 1832 to 1836”) On the Origin of Species by Means of Natural Selection, or Preservation of Favoured Races in the Struggle for Life, London 1859 Charles Darwin: On the Origin of Species. A Facsimile of the First Edition, intr. Ernst Mayr, ­Cambridge, Mass. and London 1964 The Descent of Man, and Selection in Relation to Sex, London 1871 The Expression of Emotions in Man and Animals, London 1872 Darwin und Darwinismus. Eine Ausstellung zur Kultur- and Naturgeschichte, exh. cat., eds. BodoMichael Baummunk and Jürgen Riess, Deutsches Hygiene-Museum, Dresden 1994; Berlin 1994 Darwin: Kunst und die Suche nach den Ursprungen / Darwin and the Search for Origins, exh. cat., eds. Pamela Kort and Max Hollein; Schirn Kunsthalle, Frankfurt am Main 2009; Cologne 2009 Davies, Paul Sheldon, Subjects of the World: Darwin’s Rhetoric and the Study of Agency in Nature, Chicago and London 2009 Davies, Roy, The Darwin Conspiracy: Origins of a Scientific Crime , London 2008 Davies, Roy, “1 July 1858: what Wallace knew; what Lyell thought he knew; what both he and Hooker took on trust; and what Charles Darwin never told them”, Biological Journal of the Linnean Society, CIX / 3 (June 2013), pp. 725–36 Dawkins, Richard, The Greatest Show on Earth: The Evidence for Evolution, New York and London 2009

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Voigts-Virchova, Eckart, Barbara Schaff and Monika Pietrzak-Franger, eds., Reflecting on ­Darwin, Burlington, Vt. 2014 Vollmer, Carl Gottfried Wilhelm (writing as W.F.A. Zimmermann), Die Wunder der Urwelt, Berlin (1855) 1881 Voss, Julia, “Augenflecken und Argusaugen: Zur Bildlichkeit der Evolutionstheorie”, in: Bild­ welten des Wissens. Kunsthistorisches Jahrbuch für Bildkritik, I / 2 (2003), pp. 75–85 Voss, Julia, Darwins Diagramme: Bilder von der Entdeckung der Unordnung, Berlin 2003 Voss, Julia, “Depictions as surrogates for places: from Wallace’s biogeography to Koch’s dioramas”, Philosophy & Geography, VI / 1 (2003), pp. 59–81 Voss, Julia, “One Long Argument. Die Darwinismus-Debatte im Bild”. Doctoral dissertation, ­Humboldt-Universität zu Berlin 2005 Voss, Julia, Darwins Bilder: Ansichten der Evolutionstheorie 1837 bis 1874, Frankfurt am Main 2007; Eng. trans. Lori Lantz, Darwin’s Pictures: views of evolutionary theory, 1837–1874, New Haven and London 2010 Wallace, Alfred Russel, “On the Law which has regulated the Introduction of New Species”, The Annals and Magazine of Natural History, second series, XVI / 93 (September 1855), pp. 184–96 Wallace, Alfred Russel, “Attempts at a Natural Arrangement of Birds”, The Annals and Magazine of Natural History, second series, XVIII / 105 (September 1856), pp. 193–216 Wallace, Alfred Russel, “On the Tendency of Varieties to Depart Indefinitely from the Original Type”, Journal of the Proceedings of the Linnean Society of London: Zoology, III / 9 (20 August 1858), pp. 53–62 Wallace, Alfred Russel, The Malay Archipelago: The Land of the Orang-Utan and the Bird of ­Paradise. A Narrative of Travel, with Studies of Man and Nature, 2 vols., London 1869 Wallace, Alfred Russel, Contributions to the Theory of Natural Selection: A Series of Essays, London 1870 Wallace, Alfred Russel, Darwinism. An Exposition of the Theory of Natural Selection with some of its Applications, London 1889 Warren, John S., “Darwin’s missing links”, History of European Ideas, XLIII / 8 (2017), pp. 929–1001 Weigel, Siegrid, “Genealogie. Zur Ikonographie und Rhetorik einer epistemologischen Figur in der Geschichte von Kultur- und Naturwissenschaft”, in: Helmar Schwamm, ed., Bühnen des Wissens. Interferenzen zwischen Wissenschaft und Kunst, Berlin 2003, pp. 226–67 Weinert, Friedel, Copernicus, Darwin & Freud: Revolutions in the History and Philosophy of ­Science, Oxford 2009 Wendler, Reinhard, “Versuch einer Kunstgeschichte des Baumdiagramms”. MA dissertation, Humboldt-Universität zu Berlin 2003 Werner, Fritz Clemens, Wortelemente lateinisch-griechischer Fachausdrücke in den biologischen Wissenschaften, Frankfurt am Main 1972 White, Paul, “Cross-cultural encounters: the co-production of science and literature in midVictorian periodicals”, in: Roger Luckhurst and Josephine McDonagh, eds., Transactions and Encounters: Science and Culture in the Nineteenth Century, Manchester and New York 2002, pp. 75–95 Wöhler, Hans Ulrich, Texte zum Universalienstreit, Vol. 1: Vom Ausgang der Antike bis zur Frühscholastik, Berlin 1992

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Yonge, C. M., “Darwin and the Coral Reefs”, in: Samuel A. Barnett, ed., A Century of Darwin, London 1958, pp. 245–67 Young, Robert M., Darwin’s Metaphor: Nature’s Place in Victorian Culture, Cambridge 1985 Zahavi, Amotz and Avishag Zahavi, The Handicap Principle: a missing piece of Darwin’s puzzle, New York and Oxford 1997 Zimmermann, W. F. A.: see Vollmer, Carl Gottfried Wilhelm Zimmermann, Walter, Evolution. Die Geschichte ihrer Probleme und Erkenntnisse, Freiburg and Munich 1953

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Index Footnotes are listed only when the page on which they occur contains no reference to the item concerned in its main text (which will itself refer to relevant footnotes).

abstraction – Strickland’s aversion to overly abstract models imposed on nature 32–34 – in explicating diagram in “Natural Selection” manuscript, Darwin emphasises necessary schematic abbreviation 53, 56–57, Fig. 29 – cursory sketch of seaweed (1843), as favoured “simile”, resembling abstract essence of Patagonian specimen (Fig. 1) 88–89, Fig. 45 Agassiz, Jean Louis Rodolphe (1807–73) – as co-author of Outlines of Comparative Physiology (1848) 39–42 – influence of his circular model on Darwin’s circle segment drawings 39–42, Fig. 21; 44 Alberti, Leon Battista (1404–72) on naturally occurring objects as “works of art” 4 algae: see coralline algae Amphiroa Orbignyana (Fig. 1): see Bossea / Bossiella orbigniana aquaria (coral and the 19th-century vogue for) 79 Aquinas, Thomas (1224–74) 10, Fig. 3 arbor porphyriana 10–11, Fig. 3 arbre généalogique, devised by Duchesne for strawberry (1764) 12 Ariel, in The Tempest, invokes image of coral ix; 4; 76 “aura”, assumed by items in Natural History collections 3–4

Baer, Karl Ernst von (1792–1876) and theory of archetypes 26 Bahia, Brazil, impact of its tropical landscape on Darwin (1832) 82 Barry, Martin (1802–55) – transmission of Karl Ernst von Baer’s theory of “archetypes” 26 – Tree of Animal Development (1837) 26, Fig. 15 Beagle, H.M.S.: British Admiralty-sponsored hydrographic survey expedition of 1831–36 1; 7; 23; 81–82 Berlin, Humboldt University ix Bible – Book of Genesis on Creation as single Divine Act 7–10, 11, 13 – Lamarck contrives to square his theory with biblical account 7–10, 15 – Darwin implicitly contradicts biblical account of Creation 18 “big book on species” (as Darwin’s informal working title for “Natural Selection” manuscript of 1856–58) 51 Biology, parallels with Art History during 19th century 5 birds – Strickland’s “map” of “natural affinities” (1840) 35–36, Fig. 19 – Wallace’s diagram of affinities between bird families (1856) 46–47, Fig. 25. Bolabola (Bora-Bora) in south-western Pacific Ocean 81, Fig. 43

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Bonnet, Charles (1720 –93), “scale of nature” (1764–65) 11–12, 13–14, Fig. 4 Bonpland, Aimé (1773–1853), as Humboldt’s travelling companion 83 n23 Bossea / Bossiella orbigniana, coralline alga – among specimens amassed by Darwin (who believed it to be a small coral) 1–3, Fig. 1 – Darwin appears to have derived form of 1859 diagram of “natural selection” from that of his specimen found in 1834 69–71, Fig. 36 British Association for the Advancement of Science (and Strickland in 1840) 35 Bronn, Georg Heinrich (1800–62) – many ideas similar to those of Darwin 52 – model of emergence of living organisms (1858) 52–53, Fig. 27 Carpenter, William Benjamin (1813–85), is consulted by Darwin (1846) on an adept draughtsman 71–72 cartography – is applied by Strickland to modelling natural world (1840s) 34–35; 35–36, Fig.19 – Darwin’s interest in Strickland’s cartographic approach 35–36 Cassiano dal Pozzo (1558–1657) – images of coral in his Museo Cartaceo 76–77 – first owner of The Coral Fishers (Fig. 40) Cellini, Benvenuto (1500–71), coralline symbolism of Medusa’s blood in his Perseus (1558) 75–76, Fig. 39 circles – circle-based models of nature favoured by Macleay (1821) 32, Fig. 18 – circle-based models of nature deplored by Strickland and Darwin 32 circle segment drawings (Darwin, 1850s) 39, 42–46, Figs. 20, 23, 24 Copernicus, Nicolaus (1473–1543), polymath now chiefly celebrated as mathematician, astononer and proponent of Heliocentrism, comparable to Darwin in daring to challenge received ideas 18

coral(s) – invoked by Ariel in The Tempest ix; 4; 76 – as typically found in a Kunst- und Wunderkammer ix; 4; 76 – Darwin assumes Patagonian find (Fig. 1) to be one of small corals 2–3 – examples of true coral 21–23, Figs. 10, 11, 12 – suitability as a model for species transmutation 20–21, 22–24, 29 – resemblance to coral in Darwin’s sketches / drawings / diagrams: 1837: Fig. 14; 1840s: Figs. 16, 17; 1850s: Fig. 24; 1859: Figs. 33, 36 – coral, rather than tree, as intended model for printed “natural selection” diagram (1859) 69–71 – privileged position in traditional Natural History: as mentioned in Ovid, Pliny; valued by collectors; symbol of metamorphosis, of “nature as an artist” 73–75, Figs. 37, 38 – token of bounty of nature and of nature at peace with mankind 76 –77, Fig. 40 – and political iconography in 16thcentury Florence, 19th-century Europe 75–76, Fig. 39; 77–79 – and 19th-century vogue for aquaria 79 – and Gustave Moreau’s Galathea (c. 1880) 79–81, Fig. 41 – Darwin’s view of coral anticipates his later ideas (1871) on “purpose” of beauty in nature 90 coralline algae 2–3, Fig. 1; 23–24, Fig. 13 coralline sketches (Darwin, 1837, 1840s) 16–18, Figs. 6, 7, 8; 24–26, Fig. 14; 27–30, Figs. 16, 17; 36 coral polypifer (colony of coral-constructing polyps) – lauded by Vollmer (1855), by Michelet (1861) 77–79 – Darwin recognises vast scale and inherent drama of its role 84 coral reefs – and Vollmer (1855), Michelet (1861) 77–79 – admired by Darwin as epitome of beauty emerging from struggle for life 81

Index



and Pyrard de Laval; as experienced by Darwin (1836); as described in Darwin’s Journal (1839) and Coral Reefs (1842) 81– 82, 83–84 – Darwin recognises in them irresistibility, yet infinite slowness, of evolution 85–86 Coral Reefs (1842) 81, Figs, 42, 43, 44 Darwin, Charles (1809–82) – coralline alga found in Patagonia (1834) and assumed, wrongly, to be coral 1–3, Fig. 1 – rejects Lamarck’s notion of “willed” transmutation; employs metaphor of “branching tree” of living beings; suspects species transmutation driven by “struggle for life” (1837) 7–10 – first coralline sketches in Notebook B; employs dotted / continuous lines to signify extinct / living forms (1837) 16–18, Figs. 6, 7, 8 – a poor draughtsman who nonetheless “thinks” through drawing 18; 70–71 – observation (1837) on “tree of life” / “coral of life” 20–21 – third coralline sketch (1837) 24–27, Fig. 14 – similar form in later sketch (1840s) 27–30, Figs. 16, 17 – friendship with and influence of Strickland (1840s); shares his dislike of overly abstract models imposed on nature 31–32 – considers Strickland among possible future editors for his work 32 – is interested by Strickland’s cartographic method for recording interrelationships in natural world 35–36 – circle segment drawings (1850s), influenced by Agassiz 39–42, 44, Figs. 20, 23, 24 – during work on 1856–58 “Natural Selection” manuscript, is persuaded, by re-reading Wallace’s 1855 essay, to readopt tree model 46–51, Fig. 26 – diagram in “Natural Selection” manuscript, with explication stressing uncertainty and requisite schematic abstraction 53, 56–57, Fig. 29



introduces (c. 1857) element of disruptive irregularity into drawn “natural selection” diagram 57–59 – is shocked to discover that Wallace (in 1858) has independently arrived at same theory of species transmutation through “natural selection” 61 – on advice of friends, embarks on shorter version of “Natural Selection” manuscript for swift publication 61–62 – publishes Origin of Species on 24 No­ vember 1859, with new version of diagram in chapter four (“natural selection”), and its explication at odds with closing tree metaphor 61–65, Fig. 33 – seems to have based part of 1859 “natural selection” diagram on abstracted form of Patagonian specimen 69–71, Fig. 36 – is deeply moved by encounter with coral reefs in Pacific and Indian Oceans (1836); writes powerfully of this in Journal (1839) and Coral Reefs (1842) 81 – is struck by slow, cumulative “constructive” capacity of coral polypifer, as observed in Keeling (Cocos) Islands 83–85, Fig. 44 – has access to vast range of sources to aid research after return from H. M. S. Beagle expedition 87– 88 – response to coral anticipates shift in later work: from “pointlessness” in nature (1859) to “purpose” of beauty in nature (1871); a “revolutionary” thinker, but equally a preserver of tradition in Natural History 89–91 Darwin, Emma, née Wedgwood (1808–96; marries Charles in 1839), and first full summary (1844) of Darwin’s theory 32 n4 Decaisne, Joseph (1807– 82), classifies (1842) Darwin’s specimen (Fig. 1) as a coralline alga 1–2 Deleuze, Gilles, and Félix Guattari, on rhizome 11 n8 diagram of “natural selection” – in “Natural Selection” manuscript of 1856 –58 52, 53–59, Fig. 29

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in Origin of Species (1859) 62–65, Fig. 33; 66–68 – partial derivation from formal essence of Darwin’s Patagonian find of 1834 (Fig. 1) 69–71, Fig. 36 disorderly profusion of nature – Strickland (1840s) on need for models to reflect this 32–34 – Darwin (1843) finds seaweed a fitting “simile” for this 88–89, Fig. 45 Donati, Vitaliano (1717– 62), “network” model for knowledge of natural world 20 dotted lines – used by Lamarck in “hybrid” ladder / tree diagram (1809) 14–15, Fig.5 – used by Darwin (1837–59) to signify extinct forms 16–18, Figs, 6, 7, 8; 39–45, Figs. 20, 23; 56, Fig. 29; 68–69, Figs. 34, 35 – used by Darwin (1859), in contrast to broken lines, in distinct sections of printed “natural selection” diagram 64–65, Fig. 33 drawings / sketches – their value as an aid to the draughtsman’s own thinking 4–5 – by Darwin: see Darwin (coralline sketches; circle segment drawings) Duchesne, Antoine Nicolas (1747–1827), arbre généalogique for strawberry (1764) 12 Ekkehard von Aura (1050–1126), genealogical tree Fig. 2 “endless” (as a term used by Darwin) 88–90 evolution (as species transmutation): see species evolutionary change in nature, earlier percep­ tions of – notions sparked by contents of a Kunstund Wunderkammer ix – early models of (pre-Origin of Species) 10–15, Figs. 3, 4, 5; 26, Fig. 15; 35–36, Fig. 19; 39–42, Fig. 21; 46–47, Fig. 25; 52–53, Fig. 27 extinction: see species FitzRoy, Capt. Robert (1805–65) – in command of H. M. S. Beagle on surveying voyage of 1831–36 1



accompanies Darwin (1836) in exploring Keeling (Cocos) Islands 84 n25, Fig. 44 Florence – Loggia dei Lanzi (Cellini’s Perseus) 75– 76, Fig. 39 – Palazzo Vecchio (Morandini’s Allegory of Water in studiolo) 75, Fig. 38 Galathea, by Gustav Moreau (c. 1880) 79– 81, Fig. 41 Gould, A. A. (Augustus Addison) (1805–66), as co-author with Agassiz 39–40 Gray, Asa (1810–88), pre-eminent American botanist, is sent (1857) extract of Darwin’s “Natural Selection” manuscript, as his “notion on the means by which nature makes her species” 50 Harvey, William Henry (1811–66) – Keeper of Herbarium at Trinity College Dublin, to which Darwin donates some of his collection and related n2 – registers (1847) Darwin’s Patagonian specimen (Fig, 1) as among the “Corallinae” 2 Henslow, John Stevens (1796 –1861) – and Darwin (in 1834) on indeterminate status of “Corallinae” 2–3 n7 – gives Darwin (in 1831) Vol. I of Humboldt’s account of his Central and South American journey 83 n23 Hogarth, William (1697–1764), on “variety” in The Analysis of Beauty (1753) 33–34 Hooker, Joseph Dalton (1817–1911) – close friend of Darwin 2 – as talented draughtsman 18 – co-organises (1858) joint DarwinWallace presentation at Linnean Society 61–62 – Darwin enthuses (1846) on S. W. Leonard as draughtsman 72 – Darwin claims (1844) that telling Hooker of his belief in species transmutation is “like confessing a murder” 86 n30 Hooker, William Jackson (1785–1865), and Herbarium Hookerianum 2 Horae Entomologicae (Macleay, 1819, 1820) and circle-based models 32, Fig. 18

Index

Humboldt, Alexander von (1769–1859) – travels of 1799–1804 in Central and South America 82 – strong influence on Darwin of sub­ sequent account of these, read in English translation, as Personal Narrative 82–83 Indian Ocean (in spring 1836) – Darwin’s only direct encounter with a coral atoll 83–84 – Darwin’s account of this experience in his diary and in his Journal (1839); its influence also on text and images in his Coral Reefs (1842) 83–84, Fig. 44 Keeling (Cocos) Islands, Indian Ocean, as explored by Darwin and surveyed by FitzRoy in 1836 84, Fig. 44 Kunst- und Wunderkammer (in 16th and 17th centuries) as context for thoughtprovoking encounters with coral ix Lamarck, Jean Baptiste Pierre Antoine de Monet (1744–1829) – notion of “willed” transmutation (“le transformisme”) rejected (1837) by Darwin 7–10; 20–22 – diagram of emergence of diverse animal groups published in his Philosophie zoologique (1809) 13–15, Fig. 5 Leipzig, Museum Richterianum 74–75, Fig. 37 Leonard, Samuel William, draughtsman, works with Darwin (1846) 72 Lindley, John (1799–1865) – publishes Natural System of Botany (1830; 1836) 36 – Darwin remarks on its need for diagrams 36 London – Linnean Society 61–62 – Natural History Museum 2 – Royal Botanic Gardens, Kew 2 Lully, Raymond (Ramon Llull) (1232–1316) 11 Lyell, Charles (1797–1875) – admires Wallace’s essay of 1855 61 – co-organises joint Darwin-Wallace presentation at Linnean Society in 1858 61–62

Macleay, William Sharp (1792–1865) – as proponent of circular models 32–33 – uses these in Horae Entomologicae (1819, 1821) 32, Fig. 18 macro- / micro-evolutionary change in Darwin’s “natural selection” diagram (1859) 68 maps: see cartography Medici, Francesco I de’ (1541–87) 75, Fig. 38 Medusa, and coralline symbolism of her blood in Cellini’s Perseus 75–76, Fig. 39 metamorphosis (and coral) – coral traditionally seen as epitomising this; as featured in Ovid’s Metamorphoses 73 – Darwin’s ability to fuse such a view of coral with insights gained through experience and acquired scientific knowledge 87–88 metaphors / models in Biology, and Strickland 31, 32, 33 Michelet, Jules (1796–1874), on coral in La Mer (1861) – notes its indeterminate status 74 – lauds “constructive” capacity of coral polypifer 78–79 – claims perfect allegory of Nature is a figure enthroned in coral 79–81, Fig. 41 – compared with Darwin (in late 1850s) as regards stance towards tradition of Natural History 86 models (pre-Origin of Species) of evolutionary change 10–15, Figs. 3, 4, 5; 26, Fig. 15; 35–36, Fig. 19; 39–42, Fig. 21; 46–47, Fig. 25; 52–53, Fig. 27 Morandini, Francesco (1544–97), coral featured in Allegory of Water 75, Fig. 38 Moreau, Gustave (1826–98), Galathea (c. 1880) with eponymous figure both crowned and enthroned in coral 79–81, Fig. 41 Murray, John, publisher of Origin of Species (1859) 63 n4 Museum Richterianum: see Leipzig Natural History, Darwin and its tradition ix–x; 86; 90 “natural selection” (as process driving species transmutation / evolution)

115

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Index

Darwin’s drawn diagram (c. 1857) 53– 60, Fig. 29 – Darwin’s printed diagram (1859) 62– 65, Fig. 33 “Natural Selection”, Darwin’s manuscript of (1856–58) – sends (1857) extract to Asa Gray 50 – diagram of “natural selection” in its sixth chapter 53–60, Fig. 29 – is urged by friends (1858) to publish rapidly a summary of its chief arguments 61 Notebook B (Darwin’s sketches and observations of 1837) 7–10; 16–18, Fig. 6;

Pliny the Elder (23–79), on coral in Natural History 73 “pointlessness” / “purpose” in Darwin’s perception of nature 89–90. Porphyry (233–304), and arbor porphyriana as model of natural world and related conceptual entities 10–11, 13, Fig. 3 Port Desire (Puerto Deseado), site of Darwin’s Patagonian find (Fig. 1) 1 Pyrard de Laval, François (1570–1620), as quoted by Darwin in Coral Reefs (1842) 81–82

Orbigny, Alcide d’ (1802–57) 1–2, Fig. 1 Origin of Species (1859) – derived from “Natural Selection” manuscript (1856–58) 62 – chapter four includes key “natural selection” diagram 62, Fig. 33 – explication of diagram 65 – explication of diagram (Fig. 33) at odds with tree metaphor at close of chapter four 66–69 – “endless forms” invoked in book’s rousing final phrase 89 ornithology – and Strickland 35–36, Fig. 19 – and Swainson 32 – and Wallace 46–47, Fig. 25 Ovid (Publius Ovidius Naso) (43 BC –AD 17), on coral in Book IV of Metamor­ phoses 73

Royal Botanic Gardens, Kew: see London Rüling, Johann Philipp (1741–1803) 12



Pacific Ocean (in autumn 1835) – coral reefs 81–82 – described in Darwin’s Coral Reefs (1842) 83–84, Figs. 42, 43 Pallas, Peter Simon (1741–1801), organisation of living beings envisaged in model of double-trunked tree (1787) 11–13 Petrarch (Francesco Petrarca) (1304–74) 19 Petrarch Master (fl. 1500 / 1530), satirical “tree of life” 19–20, Fig. 9 Philippi, Rudolf (1808–1904) 2 n6 Pietro da Cortona (Pietro Berrettini) (1596– 1669), The Coral Fishers 76–77, Fig. 40

Quinarian system (based on groups of five) and Macleay 32–33 n9

seaweed, favoured over tree as “simile” (1843) 88–89, Fig. 45 Shakespeare, William (1564 –1616), coral invoked by Ariel in The Tempest ix; 4; 76 species (survival / variation / extinction / transmutation) – as understood by Darwin ix; 7–10; 16–18, Fig. 6; 18–19, 20–21, 22–23, 24–30, Figs. 14, 16; 39–46, Figs. 20, 23, 24; 50, Fig. 26; 53–59, Fig. 29; 61–65, Fig. 33; 67–68; 87–89 – as understood by Wallace 46–49, Figs. 25, 26 Strickland, Hugh Edwin (1811–53) – friendship with Darwin; deplores imposition of over-abstract models on nature; and metaphors in Biology 31–34 – cartographic method of great interest to Darwin 34–35 – maps “natural affinities” among birds 35–36, Fig. 19 survival: see species Swainson, William (1781–1855) 32 time, passage of – in coralline sketch (Fig. 14) of 1837 36 – in circle segment drawings (1850s) 39–46, Figs. 20, 23, 24

Index



in drawn (c. 1857) and printed (1859) diagram of “natural selection” 53–57, Fig. 29; 62–65, Fig. 33 transmutation: see species tree (model / metaphor / simile) – Darwin employs metaphor of “branch­ ing tree” in Notebook B (1837) 10 – “tree of life” and tradition of genealogical trees 10, Fig. 2 – Neo-Platonist arbor porphyriana 10–11, Fig. 3 – Pallas and double-trunked tree (1787) 11–13 – Duchesne’s arbre généalogique (1764) for strawberry 12 – Lamarck’s hybrid of tree and ladder models (1809) 13–15, Fig. 5 – satirical “tree of life” (1532) 19–20, Fig. 9 – Darwin’s “tree of life” / “coral of life” observation in Notebook B 20–21 – Barry’s Tree of Animal Development (1837) 26, Fig.15 – Strickland’s favours “disorderly” bush model over tree model 34 – Wallace’s minimal tree model of affinities between bird families (1856) 46– 47, Fig. 25 – Wallace’s use of tree metaphor for intellectual difficulty of charting evolutionary change (1855) 47– 49 – Darwin influenced (c. 1857) by rereading Wallace to re-adopt tree model 48–51 – Bronn’s tree model of living organisms (1858) 52–53, Fig. 27 – “natural selection” diagram in Origin of Species (1859) largely misunderstood as modelled on a tree 66; 68–69 – Darwin’s explication of this diagram at odds with fourth chapter’s tree metaphor 66–68 – influence on readers of rousing tree metaphor at close of fourth chapter 67–68 – Darwin recognises value of tree metaphor as vivid and persuasive expositional device 68 – Darwin (in 1843) declares “a tree not good simile” 88

Tsarskoye Selo: Catherine Palace, The Coral Fishers Fig. 40 variation: see species Vollmer, Carl Gottfried Wilhelm (1797– 1864), on coral in Die Wunder der Urwelt (1855) 77–78 Wallace, Alfred Russel (1823–1913) – essay (1856) on “affinities” between bird families 46 – 47, Fig. 25 – essay (1855) using tree metaphor for intellectual difficulty of charting evolutionary change 47– 48 – adventurous thinking and eloquence of expression both impress and trouble Darwin 48–51 – notion of “antitypes” as possible influence (c. 1857) on Darwin’s drawn “natural selection” diagram 56, Fig. 29 – essay (1858) independently arrives at Darwin’s own theory of species transmutation through a process of “natural selection” 61 – joint Darwin-Wallace presentation at Linnean Society, London on 1 July 1858 61–62 – publishes own “natural selection” essay in 1858 and again in 1870 62 n2 Waterhouse, George Robert (1810–88), letter from Darwin (1843) attacking Macleay’s circle-based models 32 n9 West, William, medical and scientific illustrator; supplies lithographic version of Darwin’s diagram of “natural selection” for inclusion in Origin of Species (1859) 63 n4 Whitsunday Island (Pinaki), in south-western Pacific Ocean 81, Fig. 42 Williams, Helen Maria (1761–1827) – translates Humboldt’s account of his travels, the Relation historique, as Personal Narrative (1814–29) 83 n23 – Darwin is deeply influenced by reading Humboldt in this translation 83

117

Picture Credits Fig. 1 London, Natural History Museum. Fig. 2 reproduced from: Christiane Klapisch-Zuber, Stammbaüme. Eine illustrierte Geschichte der Ahnenkunde, Munich 2004. Fig.3 reproduced from: De arbore, ed. Angela Adriana Cavarra, Rome 1991. Fig.4 from: Charles Bonnet, Contemplation de la Nature, Vol. I, Amsterdam 1764. Fig. 5 from: Jean Baptiste Lamarck, Philosophie zoologique, Vol. II, Paris 1809. Figs. 6, 7, 8 Cambridge University Library. Fig. 9 reproduced from: Genealogie als Denkform in Mittelalter und Früher Neuzeit, Tübingen 2000. Figs. 10, 11, 12 Berlin, Museum für Naturkunde. Fig. 13 London, Natural History Museum. Fig. 14 Cambridge University Library. Fig. 15 reproduced from: Julia Voss, Darwins Diagramme. Bilder von der Entdeckung der Unordnung, Berlin 2003. Fig. 16 reproduced from: Studies in History and Philosophy of Science, XXVII / 1 (1996). Figs. 17, 18 Cambridge University Library. Fig. 19 from: William Sharp Macleay, Horae Entomologicae, Vol. I, part 2, London 1821. Figs. 20, 22 Cambridge University Library. Fig. 21 from: Louis Agassiz and A.A.Gould, Outlines of Physiology, London and Boston 1851. Fig. 23 Cambridge University Library. Fig. 24 Cambridge University Library. Fig. 25 from: Annals and Magazine of Natural History, sixth series XVIII / 105 (September 1856). Fig. 26 Cambridge University Library. Fig. 27 from: Heinrich Georg Bronn, EntwicklungsGesetze der organischen Welt, Stuttgart 1858. Fig. 28 reproduced from: R.C.Stauffer, ed., Charles Darwin’s “Natural Selection”, Cambridge 1975. Figs. 29, 30, 31 Cambridge University Library. Fig. 32 Cambridge University Library. Figs. 33, 34, 35 from: Charles Darwin, On the Origin of Species, London 1859. Fig. 36 detail of Fig. 1 superimposed upon detail of Fig. 33. Fig. 37 from: Museum Richterianum, Leipzig 1745. Fig. 38 reproduced from: Mario Bucci, Lo studiolo di Francesco I, Florence 1965. Fig. 39 photograph: Horst Bredekamp. Fig. 40 reproduced from: Gazette des Beaux-Arts, sixth series, CXXXVIII / 2 (autumn and winter 2001). Fig. 41 Paris, Musée d’Orsay. Figs. 42, 43, 44 from: Charles Darwin, The Structure and Distribution of Coral Reefs, London 1842. Fig. 45 Cambridge University Library