Lamarck: The Birth of Biology
Transcription
Lamarck: The Birth of Biology
Lamarck: The Birth of Biology Author(s): Frans A. Stafleu Reviewed work(s): Source: Taxon, Vol. 20, No. 4 (Aug., 1971), pp. 397-442 Published by: International Association for Plant Taxonomy (IAPT) Stable URL: http://www.jstor.org/stable/1218244 . Accessed: 24/12/2012 16:29 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. . International Association for Plant Taxonomy (IAPT) is collaborating with JSTOR to digitize, preserve and extend access to Taxon. http://www.jstor.org This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions TAXON 20(4): 397-442. AUGUST 1971 LAMARCK:THE BIRTH OF BIOLOGY Frans A. Stafleu "A long blind patience, such was his genius of the Universe" (Sainte Beuve) Summary A review of the development of Lamarck'sideas on biological systematibswith special reference to the origin and development of his concept of organic evolution. Lamarck's development towards biological systematics is traced through his early botanical and geological writings and related to the gradual change in his scientific outlook from a static and essentialist view of nature towards a dynamic and positivist concept of the life sciences as a special discipline. This development is seen against the background of the general cultural developments during the course of his life. From classic to romantic . Life and works . . . Early botanical work (1779-1800) . .. .. .. From gradation to descent (1800-1803) .. Biological philosophy and classification (1803-1809) . Lamarckiansystematics in the Philosophie zoologique (1809) Last years: an aftermath too long (1809-1829) . . The conspiracy of silence . . . . . . . . . . Appendix . . . . . . . . . . . 397 399 404 416 420 423 434 436 439 From 'classic' to 'romantic' Of the three leaders of the botany of the Enlightenment,all three stimulated by the exuberant flight of Buffon's scientific fancy, Adanson, Antoine-Laurent de Jussieu and Lamarck, it was undoubtedly the third who made the deepest impression on biology in general. Lamarck, more than any other French biologist, exemplified what Lovejoy (1936) has called "the temporalizing of the, great chain of being." Before Lamarck botanical and zoological thought was dominated by a platonic tradition with an ambivalent approach towards nature: 'nature is everywhere the same;' but also: 'nature's diversity aims at a maximum.' This paradox rests on the Platonic contingency of the two gods: the immutable, self-sufficient god or demiurg of self-contained perfection versus the dynamic and creative demiurg which manifests itself in time. A similar contingency, not fully covering this fundamental paradox of the demiurges, is the conflict in natural history between an a priori static concept of nature and a dynamic view based on experience and observation, ready to admit change and development. Another way to formulate this fundamental split is to contrast the classical attitude of the mind with the romantic: universality and continuity on the one hand, diversity and development on the other. In the course of the eighteenth century such biologists as Buffon, Haller, Erasmus, Darwin and Bonnet - partly influenced by and partly causing, themselves, a shift in society from classical 'right' to romantic 'left' - set in motion the development towards a dynamic and 'romantic' science. The cosmic order, especially its biological part, was no longer considered to be essentially static, with all recognized change being part of an essentially permanent pattern. 397 AUGUST1971 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions This old pattern was one of an infinitely static diversity - the chain of being alien to time. The new view stressed increasing diversification, development in time, creativity rather than creation. Man's task was to imitate god, therefore he should be creative, not only in art but also in science; he had to develop his individual character rather than make it conform to general standards. This emphasis on creativity, individualism and diversity is the essence of the romantic movement in science and art. In science it is unusual to employ this term 'romantic' for the breakthrough towards empiricism and positivism: the word may well serve, however, to indicate the intimate link between science, art and society. In society we see a breakthrough towards 'progress,' thanks first to the Enlightenment and later to actual revolution; in the arts romanticism develops through the change from 'form' to 'contents;' in philosophy the emphasis shifts from mind to life, from noumenal ego (Kant) to temporal ego; science progresses from essentialism via temporalizing nominalism to pure empiricism: briefly, a shift from uniformitarianism to diversitarianism. Buffon introduced the time element in biology and drew attention to biological change and variation, even though his mind stayed in many other respects with the processes of simplification and standardization which were so characteristic of the early Enlightenment. Adanson introduced a strong impetus of empiricism and inductive research into an essentially still static concept of the order of life, but he also failed to dissociate himself fully from the uniformitarian concept of nature: he tried to establish 'the' rather than 'a' natural system. A.L. Jussieu, even more than Adanson, refined empirical research, but he framed the results within an essentially static concept of life. For Jussieu the time element was irrelevant where botany was concerned: all temporal processes were ultimately cyclical. Buffon alone had daringly suggested the possibility of an open-end concept of life in opposition to that of a closed circuit. Lamarck was perhaps the most involved personality among these four French biologists. Only he had a really 'cosmic' approach, because he gave equal attention to physics, chemistry, the earth sciences, and biology. The development of his thinking involved a more fundamental change with respect to the age-old principle of plenitude than was the case with the others. In various respects Lamarck's world of thought remained part of the past; in several fundamental issues, however, his thoughts were spearheads toward the future. Even though elements of truly evolutionary thinking can be found in the writings of some of his predecessors and contemporaries (e.g. Buffon and Erasmus Darwin) Lamarck was the first to formulate a full-fledged transformist theory. He did this on the basis of a division in principle between the world of living beings and that of 'mineralogy.' This division was based, as we shall see below, on an essentially erroneous chemical theory, but his main general conclusions have proved to be correct, although perhaps not for the reasons he gave. His theory of organic evolution was based on assumptions which have not been proven: even so, the principle of an open-ended development of life, now called evolution, is generally accepted. Lamarck started out with an essentially static taxonomic theory based on the scala naturae, but was the first to state explicitly that it could - in amended form - be taken as the mirror of evolutionary development. Lamarck's own contributions toward taxonomy remained traditional to a great extent; the advanced elements of his cosmology, on the contrary, cleared the way towards evolutionary thinking. He was not to witness the final victory of evolutionary thought; for this he was born half a century too early. 20 VOLUME TAXON 398 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions Life and works Buffon's life is well known and richly documented, although the definitive biographer has not yet presented himself; Adanson's place in the history of science has been the subject of a small number of fairly recent publications (see e.g. Nicolas 1963, Stafleu 1963, 1967); Antoine-Laurent de Jussieu has been relatively modestly treated by the historians of biology. For Lamarck, however, there is no lack of documentation and comment. The virtual - if not actual - founder of transformism has received ample and diversified biographical attention in print e.g. by Packard (1901), Landrieu (1909), Kuehner (1913), and Tschulok (1937) and more recently for instance Gillispie (1956), to mention only a few. A brief biographical sketch may therefore suffice here. Jean Baptiste Antoine Pierre Monnet de Lamarck was born on 1 August 1744 at Bazentin-le-Petit, a small village in Picardy in what is now the Department of the Somme. Lamarck's father was listed in the register of birth at Bazentin as "Jacque[s] Philippe de Monet, chevalier, seigneur des Bazentin grand et petit." De Monet was the patronym; his domanial name was de Lamarck or Lamark. The five main periods of Lamarck'slife are (Tschulok 1937): 1. 1744-1761, childhood at home and education at the Jesuit college of Amiens, 2. 1761--1768, army career, 3. 1768-1778, bank employee in Paris until the publication of the Flcre francoise, 4. 1778-1793, botanist at the Jardin des plantes, 5. 1793-1829, professor of zoology at the Museum d'histoire naturelle. Lamarck, the youngest of eleven children, was originally meant to follow an ecclesiastical career. At the age of seventeen, however, he joined the army and took part in one of the seven year's war's battles in Germany (near Lippstadt); then he spent seven years in garrison at Toulon and Monaco. He was pensioned off because of a minor physical disability and settled at Paris in the course of 1768. He accepted several minor jobs which enabled him to study medicine and botany, the latter mainly with Bernard de Jussieu. There are indications that he met Jean Jacques Rousseau, but on the whole little is known of his activities during these years. The main result, however, was a higly original work, the Flore franqoise ou description succincte de toutes les plantes qui croissent naturellement en France, dispose'e selon une nouvelle methode The book, 3 volumes, dated 1778, and signed "Par M. le ched'analyse..." valier de Lamarck" appeared in March 1779 at the Imprimerie Royale. Lamarck had offered it for publication to the Academy of sciences, which recommended it for publication. The book was in several ways a 'first'; it was the first modern-type flora of France, it was written in the vernacular (in agreement with the Enlightenment ideals of wide diffussion of knowledge) and it presented a simple and effective disposition of all species in the form of dichotomous or nearly dichotomous keys. It is likely that Lamarck had by that time been discovered by Buffon and that the latter was instrumental in having the book printed at the expense of the state, with the royalties coming to the author. Such additional income must have been very welcome because Lamarck lived on a small army pension supplemented by a modest income from his share of his parents' estate. Further financial relief came in the same year, 1779, when he was appointed 'adjoint' member of the Academie des Sciences. The competition was between the physician Descemet and Lamarck; the former had the support of the other botanists, Jussieu and Adanson, and was listed as number AUGUST 1971 399 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions one, but the king followed Buffon's advice and appointed Lamarck. This unusual procedureshows the influence of the intendant du lardin du Roi, Buffon. The latter then requested Lamarck to accompany his son on a grand tour through Europe for a visit to Holland, Germany, Bohemia, Austria and Hungary; a tour on which he made the acquaintance of renowned botanists such as Gleditsch, Murray and Jacquin. Back in France in April 1782 Lamarck found himself in slightly easier circumstances because of his salary (pension) as a member of the Academy. No place being available, however, for a fully salaried position at the Jardin du Roi, Lamarck set out to compile his most voluminous botanical work: the botany volumes of the Encyclopedie methodique, the continuation "par une societe de gens de lettres, de savans et d'artistes" of the original Encyclopedie by Diderot and d'Alembert. The Encyclopedie methodique was the apogee of the encyclopaedic movement of the Enlightenment.Not less than 196 'Tomes' (186 vols.) were published between 1782 and 1832 dealing with zoology, anatomy, botany, agriculture, chemistry, social sciences, history, arts, geography, games, etc. Among the first volumes published in 1782 were part of 'jurisprudence' and a general zoological volume by Daubenton. The moving force behind this almost incredible outburst of synthetic scientific creativity was the publisher CharlesJoseph Panckoucke (1736-1798) whose father Andre (17001753) had had a publishing firm in Lille. Pancoucke published e.g. the Mercure de France, the supplement of the original Encyclopedie, the collected works of Voltaire and of Buffon and the famous Moniteur universel. The original plan was to publish the Encyclopedie methodique between 1782 and 1787 with 53 volumes of text. It appeared in 'livraisons' of several volumes at the same time. The sixth livraison, published on 2 Dec. 1783, contained Lamarck's volume on botany, 'tome' 2(2) of the arts et metiers mdchaniques, and the second volume of plates. The botany volumes of the Encyclopedie, published between 1783 and 1823, first by Lamarck and later by Poiret and other collaborators, constitutedthe most detailed systematic survey of the plant kingdom of its time. The work on the Encyclopedie took almost all of Lamarck's time between 1782 and 1788, the year in which he was appointedto the newly-createdposition of keeper of the herbarium of the Jardin du Roi, with a salary of 1000 francs and the official title botaniste du roi avec le soin et la garde des herbiers. The herbarium of the Jardin du Roi was at that time insignificant. All botanists, whether attached to the Jardin du Roi or not, had their private herbaria which were often infinitely richer than the paltry holdings of the royal natural history cabinet. The botanists on the staff were allowed to build up private collections even by means of material collected on state paid collecting trips. The Jardin du Roi dates from 1597, although functionally only from 1635 when its third curator, Guy de la Brosse, really set things moving. The first herbaria of any size to be acquired by the garden were those of Tournefort (1708) and Sebastien Vaillant (1669-1722). This was about all Lamarck found to 'curate.' In addition there were other collections such as those of Commersonand Dombey, but they were on loan to Jussieu and L'Heritier for publication. Lamarck'sappointment was therefore a sinecure: under the existing system he could do little to increase the size of the collections. For this France needed a revolution. Lamarck's life, as that of most people, was complicated during the years 1789-1793 by the general social and political events. At first the royal intendant for the garden, La Billarderie, tried to suppress Lamarck'spost as keeper of the herbarium (Hamy 1893). Lamarck reacted (1790) by publishing two pamphlets, of which the first is a MImoire sur les cabinets d'histoire naturelle, 400 TAXON This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions VOLUME 20 et particulierementsur celui du Jardin des plantes.. ." In this treatise Lamarck unfolds his ideas on the reorganization of the garden, emphasizing its public r^le and duties. The use of the name Jardin des plantes in print - so far this had been a more or less colloquial synonym for the official Jardin du Roi was in open defiance of the royal set-up and had all the flavour of the revolution. The staff held its own meetings and addressed itself to the Convention. After a curious interregnumbetween 1791 and 1793, when Bernardin de Saint-Pierre was the last intendant of the old institution, the Convention authorised the professors on 10 June 1793 to constitute an assembly and to nominate a director and other officers. The Museum d'histoire naturelle decrte' par la Convention nationale le 10 juin 1793 came into being at the assembly of 14 June. Daubenton was chosen as director; Lamarck and Fourcroy were commissioners for the formation of a museum library; Jussieu and Desfontaines were charged with the building up of a herbarium.The libraries of convents and of many aristocrats were to be nationalized, and the revolutionary armies were instructed to confiscate the herbaria found in 'liberated' countries. The professorships were newly divided, and Lamarck was appointed 'professor of zoology, of insects, of worms and microscopic animals.' Even though he had been mainly a botanist before this 49th year of his life, Lamarck came to zoology not quite unprepared. His knowledge of taxonomic methodology obtained in botany provided an excellent background for zoological work as well. Furthermore,Lamarck had been an enthusiastic conchologist for many years; he was an avid collector and had been remarkably successful in building up a natural history cabinet of his own. His herbarium was one of the largest of his time and contained material from all parts of the world. Lamarck'sshell collection was a match to this herbarium, and the shift from botany to zoology cannot really have been such a hardship as some of Lamarck'sbiographers have supposed. The appointment to the chair of invertebrate zoology was an important reason why Lamarck turned away from botany after 1793. He had done his share by publishing the Flore franqoise, the greater part of the first three volumes of the Encyclopedie, the first five volumes of the Tableau encyclopedique (900 plates with descriptive text) and a number of smaller publications in journals such as the rare Journal d'histoire naturelle of 1792. It is questionable, though, whether it was really only this appointment which made Lamarck drop botany almost entirely. His versatile mind, heavily leaning towards theoretical synthesis and towards 'the new' turned away from botany because the subject lost its challenge to him. In the years 1793-1799 Lamarck gave much but by no means all of his time towards the zoological assignment. His publications as well as contemporary testimonies show that his real interest in these years lay with his physical, chemical, geological and meteorological work. This phase in Lamarck'slife cannot be left out of account merely because the books published in these years have turned out to have had little lasting value for those particular fields, with the exception perhaps of geology; they form an essential part of Lamarck's intellectual production and cannot be dissociated from his later works. Lamarck's geological works, as reflected for instance by his Hydrogeologie (1801), with its emphasis on change and development of the earth's surface, must have made him aware of similar processes in the world of living beings, leading him almost directly to his transformist theory. His 'pyrotic' theory of chemistry was really a variant of the old phlogistian theory but fitted very well into his theory of the earth which assumed that there were two basic processes, one of synthesis through increasing complexity by life AUGUST1971 401 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions (organic nature) and another of decomposition and decreasing complexity in anorganic nature. Lamarck's theory was essentially uniformitarian but it contained several beginnings of important new departures such as the immensity of geological time, geological actualism and the transformist view of organic development. The publications of the years 1793-1799 have significant titles: 1794, Recherches sur les causes des principaux faits physiques, a manuscript dating from 1776 but for which Lamarck had previously found no publisher; 1796: Refutation de la theorie pneumatique, which attacked the advanced chemical views of people like Lavoisier (murderedduring the revolution) and proposed Lamarck's own theory of fire and chemistry; 1797: Memoires presentant les bases d'une nouvelle theorie physique et chimnique,fond&esur la consideration des molecules essentielles des composees... ; 1797: Memoires de physique et d'histoire naturelleJtablis sur des bases de raisonnementindependantde toutes theories... ; 1798: De l'influence de la lune sur l'atmosphere terrestre et les nuages; 1799: first volume of the Annuaire meteuorologique.This series was hardly interrupted by botanical or zoological publications. Some parts of the Encyclopedie written by Lamarck appeared between 1793 and 1797, but they had obviously been prepared in earlier years and were supplementedby other authors. Lamarck'slast botanical publications are the first two volumes of an Histoire naturelledes vge'taux published in 1803 (Lamarcket Mirbel 1803). A. P. de Candolle mentions in his Memoires et Souvenirs (1862) that he met Lamarck twice during these years. The first meeting in a Paris restaurant in the winter of 1796/1797 was followed by some visits at home: " ... mais comme il etait dans ce temps tout occupe de ses objections contre la theorie chimique et qu'on ne pouvait I'amener a parler de botanique, je tirai de cette connaissance moins de profit que je ne l'avais espere." In 1798 de Candolle returned to Paris and again met Lamarck, who asked him to do some articles for the Encyclopedie (e.g. Paspalum, Parthenium, Parnassia). At that time Lamarckhimself was obviously still very much interested in the publication of this great botanical compilation, although he left the work to others. De Candolle remarksthat the use of Lamarck'slibrary and herbarium was extremely useful, but again that "ce savant etait alors absorbe par ses ecrits contre la chimie moderne et par ses hypotheses relatives 'l'actionde la lune sur l'atmosphere.Quand je l'interrogeais sur la botanique il me repondait par de la chimie ou de la meteorologie qu'il savait a peine [sic]." This scant knowledge of meteorology attributed to Lamarck by the overly sceptic de Candolle produced, for instance, the very astute observation that there is a high and a low atmosphere,the first of which he called the region des meteores, or weather region (Matout 1930). Lamarck'sfirst zoological publications of any importance are purely descriptive and date from 1798 (a volume on testaceous mollusks for the Encyclopedie, his old hobby) and 1799, the year in which he proposed a new classification of the 'coquilles.' Epochal was the publication of the Systeme des animaux sans vertabres of 1801, which contained the text of the historical opening lecture of the zoological courses given on 21 Floreal, an VIII [11 Mai 1800], the lecture in which the theory of biological transformismwas first presented. From 1802 onward Lamarck produced a signal series of descriptive as well as theoreticalworks dealing mainly with evertebrateanimals and palaeontology. He continued the publication of his meteorological yearbooks, in which he tried to give long-rangeweatherforecasts. In Dec. 1801 or January 1802 there appeared 402 TAXON This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions VOLUME 20 his Hydrogeologie, which was planned as the first of three volumes that would deal with his views on 'the physics of the earth.' The subtitle of the Hydrogeologie is, translated (Carozzi 1964): "Investigations of the effects of running waters on the surface of the earth, of the reasons for the existence of an ocean basin and its gradual displacement over various parts of the earth's surface, and finally, of the influence of living organisms on that surface." This long subtitle, like a modern abstract, gives in a nutshell Lamarck's dynamic view of the origin of the crust of the earth and of the preponderantr61e played by living organisms. Lamarck states in the preface (p.p. 7, 8) (translation Carozzi 1964): "A sound physics of the earth should include all the primary considerations of the earth's atmosphere, of the characteristics and continual changes of the earth's external crust, and finally of the origin and development of living organisms [sic!]. These considerations naturally divide the physics of the earth into three essential parts, the first being a theory of the atmosphere, or Meteorology, the second, a theory of earth's external crust or Hydrogeology, and the third, a theory of living organisms, or Biology." The book on meteorology was never published, and the Biologie never appeared in this form but was in a way replaced by the Philosophie zoologique of 1809. In the Hydrogeologie we find therefore the first definition and use of the word 'biology': a theory of living organisms. Only a few months later, in the year 1802, G.R. Treviranus published the first volume of his book Biologie oder philosophie der lebenden Natur. The word 'biology' was thus born with the century, althoughthe first statementof the concept goes back to Buffon. Lamarck's ideas on the origin and development of living organisms had, in the meantime, been put forward in his opening address of the zoology course at the Museum,on 11 May 1800, and in his Systeme des Animaux sans vertebres of 1801. In all these publications Lamarck's ideas on the r1le of time and environment as the "two principal means which nature employs to bring its products into existence" were clearly expressed. The Philosophie zoologique contained much material from the opening lectures of Lamarck'szoology courses between 1800 and 1806. Lamarck explains in the 'avertissement' (p. XVIII) preceding the work that for the Philosophie zoologique he has used the "main materials which I assembled for a planned work on the living beings, under the title Biologie; a work which as far as I am concerned will not be produced [restera sans execution]." The Philosophie was followed by Lamarck's final biological work, the Histoire naturelle des animaux sans vertebres, of wich the first part, containing the general introduction, appeared in March 1815. The six subsequent volumes appeared between 1815 and 1822. In the supplement to the first volume Lamarckclearly states that so far classification had resulted in the establishment of series on the basis of agreement or difference of charac" teristics ('rapports'). However, "un pas de plus restait donc faire, c'&taitle plus important, celui meme qui pouvait le plus nous 6clairer sur les operations de la nature. 11 s'agissait seulement de reconnaltre que les portions de la serie generale que forment les objects convenablementrapproches par leur rapports, ne sont elles-memes que des portions de l'ordre de formation a l'egard de ces objects." In other words: the natural system reflects the phylogeny. The step was decisive: a new era in systematics could begin. The Histoire naturelledes animaux sans vertebreswas Lamarck'smost important synthetic survey of 'his' part of the zoological kingdom. Better known, because it was more widely distributed, is its second edition, posthumously published in 11 volumes between 1835 and 1845 and edited by G. P. Deshayes and H. Milne Edwards.The last volume of the original Histoire (1822) came out after AUGUST1971 403 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions Lamarck had become blind. Part of the book was dictated to his daughter Cornelie, who took care of her father all through his old age. During these last years Lamarck'sfinances were at their lowest ebb. His savings had disappeared, not least because of risky investments and the diminishing income from his books. In 1824 Lamarck decided to sell his herbarium. The institutionalization of herbaria was not yet so far advanced that the French state was interested in acquiring this valuable collection for the nation. The history is reminiscent of that of the Linnaean herbarium. Again another botanist from the country of origin, this time Adrien de Jussieu, was instrumentalin selling the collection abroad, no doubt in an honest effort to help the ailing and impecunious Lamarck. J.A.C. Roeper bought the herbarium (Lecomte et Leandri 1930) at the advice of Alexander von Humboldt and took it with him first to Basel (1826-1836) and then to Rostock in Northern Germany. The grand duke of Mecklenburg bought the Roeper herbarium for Rostock in 1875 but after Roeper's death (17 March 1885) Goebel decided to make the herbarium available again to science by selling it to one of the main botanical establishments. The herbarium was offered to Paris and at last acquired by the French state for the Museum. The herbarium is now at Paris, kept separately, wel arranged and available for consultation not in the least because it has also been put on microfiche (IDC no. 6207). Lamarck'slast years were sad and lonely. During the years when his eye-sight failed but had not yet completelydisappearedhe publishedthe general philosophical work Systeme analytique des connaissances de l'homme..." (1820). The last volume of the Histoire, mentioned above dictated in part after he had become completely blind, was also the last volume published by the aged scientist himself. Lamarck lived until 18 December 1829, dying after lonely dark years of invalidity, blindness and poverty. His grave was temporary: the location is known, but his remains were dispersed to anonymity at an early date. Lamarck's first book appeared in 1779, his last in 1822: forty-three years of an incredible productivity had resulted in a long series of descriptive as well as theoretical works. The descriptive publications have retained their value for biological taxonomy. In the theoretical works Lamarck revealed his genius as well as his limitations; these were so numerous and often so controversial that they were received on the whole with little enthusiasm; many of them encountered cold hostility or, worse, silence, during his life time. Even so they contained several fundamentallynew developmentsin biological thought; to mention only one: the formulation of the first scientific theory of transformism lead the way towards the later theories of evolution. This transformist theory was the logical result of Lamarck'stheoretical views on life and earth. Transformism may have had no immediate impact on the daily routine of systematics, but its long range influence was such that Lamarck's views form an essential part of the history of ideas in systematics. Early botanical work (1779-1800) In his early botanical work Lamarck shows himself a true follower of the lines of thought of Buffon, Bernard de Jussieu and Adanson. His approach to nature was twofold: he developed a purely pragmatic analytical system to facilitate identification and elaboratedthe idea of a natural system as a general expression of biosystematic knowledge. In Lamarck'sfirst work, the Flore frangoise (1778, publ. 1779) the emphasis is mainly on the first approach. The book (3 volumes) consists of a lengthy TAXON VOLUME 20 404 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions theoretical introduction followed by an 'analytical method' to name the plants occurring in France. This method is an elaborate key of the kind which has now become a routine element in any flora. It is often dichotomous and leads directly to descriptions of the species with synonymy and habitat. The species are arranged in genera, but the genera are not necessarily grouped in higher taxa. The arrangement follows from the most direct way to identify the taxa by means of the key and is therefore not in accordance with any of the existing systems or methods. The Discours preliminaire to the Flore franqoise contains Lamarck's first credo as a botanist. He remarks that all earlier systems had tried to serve two purposes but that they had failed because these purposeswere contradictory.The systems tried to paint a picture of the vegetable world, of the form-relationships between the plant species. At the same time, however, they aimed at providing an easy method for the naming and identification of plants. For the latter certain purely artificial conventions were necessary which were in flat contradiction to what Lamarckcalls la marche de la nature. It is the task of the natural method to give an overview of this master-planof nature; it is something quite different to set up a simple analysis revealing the identity of the taxa. The sole aim of the Flore franqoise is to provide such a means of identification. It is important to be clear about the meaning of the term la marche de la nature in this phase of Lamarck'sthinking. In these early years Lamarck did not question the fixity of species or the dogma of special creation. His marche, a term with dynamic overtones, is here nothing but a metaphor for the series into which living beings can be arranged in accordance with their degree of overall organization. The marche de la nature is nothing but the great chain of being; it has no componentin time. In later years Lamarckbecame convinced that this purely descriptive master-planalso revealed the order in which these living beings had been producedby nature during the immense period of geological time. This breakthrough, however, took place after 1793. The botanical works published before that year all present a picture of a timeless creation; nature is seen as an immense garden which arouses in the spectator a feeling of admiration for that "sovereign free and independent Power which manifests itself in this immense variety of beings... " All attempts to paint a picture of this universe by means of dividing lines, classes, families and genera, says Lamarck (before 1793), are nothing but an avowal of weakness by the human mind under the guise of an imposing scientific apparatus. It is clearly a follower of Buffon who speaks here: nature presents us with an immense variety and variability which defeats any attempt to depict it by means of conventional systems. The concept is essentially uniformitarian('there are no fixed dividing lines') and nominalist. Classes, families (even if called 'natural') and genera are nothing but creations of the human mind; they are subjective, because we shall always find species which are intermediarybetween whatever genera we have created for our convenience. This subjectivity is characteristic even for many of the species distinguished by botanical authors. In this case, however, we have, at any rate in principle, the possibility to grow the plants together under identical circumstances and find out whether they are then different or identical. Somewhat later Lamarck will define species as reproductive communities, assigning them an objective status. Lamarck, as Adanson and Buffon, uses the term 'system' for classifications based on a single characteristic. Methods are more flexible because they are based on a great variety of characteristics; on the whole they follow "principes moins fixes..." It would seem that a system would be more advantageous as AUGUST1971 405 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions ...... .......... ....(.. Vow-,* iLvllf;,?,?. 9AVOii 41 A "WC- b •i!!iiiiiii1%ei •iiiiii4C V4?m A.vt tv tv 4v ow c ,*fiiii 0 t cut/ ;t V.i iiAJi•! itii ~ht? ZA!iii~ .... iiii~iiiiKii •.7 t7 4,PLO lop I74~h. ~ I&J4. t4 Lei kav OLV40? ..........( ii!iiii~i1 a?5~i i:!iiiii!•iiiiill i~iii~iiiiouk VA'ttv#"JLo" L 1. owle-c ,... ir iii t....... If!lt 5 . C'O" tii 4A it 4-4CA XV-i~~ii 4?LkAAOL4 ?.iiiil FIG. 1. First page of letter by Lamarckto Pancoucke in which he discusses the working plan for the botanical volumes of the Encyclopedie methodique (30 December 1780). For a translationof this letter see the appendix to this article (p. 439). (Reproducedby courtesy of the director of the Conservatoirebotanique de Geneve). 406 TAXONVOLUME20 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions *':jijlilji jijii ijlili -VA- ij? -iiiiiiii -: .-:--.:-:::i:::::::-:i:jiiiijl.. il::::i:::: ,ii~i~ili~i~iiii::-i::::- 4L : :::::::::I :.'... ..::,:: : A. : - ? itIt ::? _ ? :: . " "iljlli :l.:iiil:iiil:i-':'i--iiiiiiiiijiiiliiii ....-::::::::::I:1::: 11::::::: i--iiii iiiiii' - -' Li:::: iii A.?vw coAig V i r:*_::::i ...... iiiiiiii-ii--i:i~ .'.ld i!-ii :iiii: -.I i ....... !iiiiii~iiii~i!•,; i~jiiiiiili -i iii~ ,.:?:: i- ii i i, :::: ::::,: ................ i i i~i:~:::::: :::::::i : ":: ::: : iii:: i :::::7~: :-::::::r: ::: T Ili~:::: --::: :-? ::::???::::::::::: ::::: ?:::::::::I ::: P~:::; ::--:t:~ ?_::_ A t ::\:::::::?::::::::::::: o:::; ::*::::::: :::::::::::::::::::::::,l;: ts:ii:: a iiil:::• 7:: : : : s; : : ?OW*Kt 04OA" .4)1044041 rwr : ?ticbow wir ts-iv,?L *I' CS, Z? tiiiiliiiiavix myo:::::ivtv : Vbliag~Qn~a~ii~" :i::liiii~iii~i ~ ~~:: Y -i ijl i~lji li: VCl vW r :y::-ii-i - :: ::ia:::-?:'':::. :?i::-:::i : _ Cft W C:l :cia . .%J::: J).:: WS:t 4v': :4A5: ::7:OI4AI: :::$4A.:::::: t4%7::PC4ItiVA4.: ::t:: ??liij:ii I:iiii:i~~~i! AL%A0 VAr10- c-oC * ~ C:/ _ll :: ::::-::::::::::i- -'c ~ ~ 4.7, U44 .r (s.sts:& v4 1 -: io: ::::::,:::--::---:-~l~:?:i::: UA::::: A,,&i-::::1::~::iV" i:::::::::::::::::::~ : .. ... ..... ....... ::.. ia'iii::?-:-? _!i:-_ji :iiiiiiii'iiiii!i .-•iiii:j~~-Li''i~ii/ ii~ji iicttiiij: ii:d:iiiiilji:iii?iiii lllli ....?:iilii :: !:::::i~c~ e ~iiiiiLiiiiiiiiiiiij 0i;ji:'lii:f~iPli ri~~~~' iiii-?-::i ravOt, l :::::::::::::::::::: ?:::: l:::: vkilliiiiii::: /XO45) t~ ili iii!ii i _iiii U .............. FIG. 2. Second page of Lamarck's letter to Pancoucke of 30 December 1780. (Reproduced by courtesy of the Director of the Conservatoire botanique de Geneve). AUGUST1971 407 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions a means to identify and name plants. The trouble is, however, that the systems are too rigid because of their emphasis on very few and a priori chosen characteristics. As a result they have to admit too many exceptions to make them really work; a fool-proof identification must remain fully flexible. Lamarck therefore advances his analytical method which keys out the plants (in principle dichotomously) on the basis of the most practical characteristics, not necessarily derived from the same organ. However, botany cannot be satisfied with identification alone. It will also be necessary to paint as true a picture as is possible of that "admirably graded chain" which seems to be presented by all plants in their form relationships. This picture, which is called the natural method, cannot also fulfil the r8le of a key. Lamarck's analytical key was a success. The taxonomic part of the Flore frangoise was indeed by far the most useful flora so far published for purposes of identification. The 'natural order' of plants was not entirely neglected either, at least not in the Discours preliminaire. The problem is how to find a means to describe such an order if one does not recognize the existence of sharp divisions. The solution is that of the series. One should first find out which taxon has to stand at the beginning because its overall organization is simplest. By general comparison this should then be followed by the taxon which shows the closest affinity to it. Lamarckeven works out a numerical system of points of resemblance which brings him closer to modern numerical taxonomy and phenetics than Adanson ever came; he even tries to weight his resemblances numerically. This method is mentioned only in passing and there are no signs that Lamarckever really establishedthe numericalrelationshipsof all his plants. A flora of France is not the place for such an attemptbut the reader is promised a Thidtre universel de botanique in which all plants will be analysed, described and arranged according to their (numerical) degree of relationship. The Thedtre never appeared. After completing the Flore frangoise and after his trip through Europe with the son of Buffon he was asked to prepare the botanical part of the great Encyclopedie mhthodiqueof Pancoucke. In the Thiatre he would have treated the plant kingdom in accordance with the 'natural order', in an Encyclopedie, however, the alphabetical arrangementwas imperative.* There are entries both for taxa and concepts. Several of the articles on the latter, such as on classes, genera, species, and botany, form together an important treatise on taxonomic methodology; the treatment of the taxa forms an almost complete Species plantarum. Because of this combination the Encyclopedie, together with the additional volumes of plates and concise diagnostic texts entitled Illustrations des genres, constitutes one of the most remarkable and extensive botanical treatises ever published. A first feature, understandable in an encyclopedia, is that the text is fully in French, like Adanson's Familles des plantes. The Encyclopedie fulfilled one of the ideals of the Enlightenment, to break the esoteric ring around science and to make it accessible to the widest possible public. Another feature is the great detail in which every species is described: full synonymy, literature references, ample descriptive morphology and information on geographical distribution. None of the earlier comprehensive treatments of the plant kingdom had ever presented so much detail. The genera are also amply described and are all attributed to the natural families. The families themselves are briefly discussed and placed in the relevant Lamarckian class, with indication of all constituent genera. These natural families are practically identical with those of Jussieu's Genera plantarum (1789); how* See figs. 1 and 2. Lamarckmentions the Th6atre here under the title flore universelle. See further below, appendix 1. 408 TAXON VOLUME20 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions ever, the first volume of the Encyclopedie came out in 1782, seven years before Jussieu's work. In his Discours preliminaire Lamarck publishes the 'Methode de M. Jussieu' (i.e., Antoine-Laurent),giving for the first time the complete list of the (French) family names. The treatments of the families contained in the first two volumes (A-Gor) have therefore precedence over those in Jussieu's own Genera plantarum. Notwithstandinghis tremendousindustry, and the rapid and efficient publication by Pancoucke, Lamarck could not terminate the work. He wrote the text for the first two volumes and part of the third (Poiret completed it); the fourth volume was still edited by him but most articles are by Desrousseaux, Poiret and Savigny. Volumes 5-8 and the supplements are by Poiret with incidental contributions by others. Poiret's achievement over the years (17921817) was formidable; unlike Lamarck, he stayed with systematic botany; his work on the Encyclopedie became his major contribution to biology. A richly illustrated synoptic review of the genera and species was published separately under the title Tableau encyclopediquedes trois regnes de la nature. Botanique,' also known as Illustration des genres. Lamarck contributed not less than 900 plates with text between 1791 and 1799; here again the work was terminated by Poiret. This synoptic treatment follows the pragmatic Linnaean artificial system (at the request of the publisher) and presents genera and species in the classical Linnaean manner. The text is in Latin and French, in parallel columns; the genera are briefly diagnosed by means of 'essential characters' and more elaborately described by 'natural characters.'The text is therefore much more old fashioned than that of the Encyclopedie: for practical purposes, and for a wider public, the Linnaean presentationwas still preferable.The most important part of the work, however, are the detailed illustrations themselves. The Illustration des genres was for a long time the most elaborate iconography of the plant kingdom. The speed with which this immense documentation was brought together did not allow the use of much living material. The artists (Audebert, Fossier, J. E. de Seve, H. J. Redoute) had to work mainly from herbarium material, often from Lamarck's own collections. All through the years 17911799 Lamarck was actively engaged in keeping the engravers at work. The Illustration was the main botanical work with which he occupied himself also after his appointmentto a zoological chair at the Museum. It is good to realize the immense amount of descriptive and critical detail assembled by Lamarckon the basis of his own and other private harbaria. Even though some of his theoretical works would seem to indicate the contrary, Lamarck was primarily an empirical scientist with a profound and diverse knowledge of plants and animals. The Discours preliminaire included in the first volume of the Encyclopedie is mainly a historical review of plant classifications much along the same lines as those of Adanson (1763). Lamarck discusses Adanson's Familles des plantes briefly, without really doing justice to the fundamentalvalue of the book. Many passages in the theoretical parts of the Encyclopidie and the Flore franqoise are very similar to passages in the Familles des plantes, without special reference to that book. Lamarckpays one handsome complimentto Adanson: "Neanmoins M. Adanson... qui s'est livre depuis long-tems avec une ardeur incroyable 'a l'tude non-seulementde toutes les parties de la Botanique, mais meme de toutes les branches de l'Histoire naturelle ... aura toujours la gloire d'avoir publie le premier des familles de Plantes determin~es dans le dessein de faire connoitre les vrais rapports des vegtaux entr'eux." Lamarck himself, however, adopts the method of the Jardin du Roi developed by the Jussieus. The Discours conAUGUST 1971 409 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions tains no methodologicaldiscussions or definitions of key concepts in systematics: for this we have to turn to the Encyclopedie itself in which, under various headings, Lamarck expresses himself on these subjects with much greater precision and clarity than in his Flore franqoise. Lamarck defines the species as follows: "en Botanique comme en Zoologie, l'espe'ceest constituee necessairement par l'ensemble des individus semblables, qui se perp&tuentles memes par la reproduction."Without constant reproduction of similar individuals one cannot speak of a true species: for this reason it is wrong to speak (as Adanson did) of species of minerals. Even before he doubts the constancy of species, therefore, Lamarck makes the fundamental distinction between living beings and non-living objects; he is no longer a natural historian but a biologist. A further statement confirms this biological species concept: it cannot be denied that the species really exist in nature. All groups of species, however, whether anitial or vegetable, are 'parfaitement artificielles.' These groupings are extremely useful arid necessary, but their origin - the human mind - must never be forgotten (article Espece, Enc. 2: 395). Lamarck adds that some botanists had doubted the existence of species, but this was because their delimitation was at fault: species can be recognized by clear characteristics ('caracteres tranchans') "always confirmed by constancy in reproduction." The multiplication of 'bad' species common in his day could only occur because botanists lost sight of this fundamental requirement. Simple environmental varieties can always be recognized as such by cultivation under identical circumstances. From the above it follows that genera are established for convenience: Lamarck'sattitude is purely nominalist and pragmatic (Enc. 2: 630). The main criteria used when grouping species into genera are derived from the characters of the fructification. Species with an almost identical fructification will usually constitute a single genus; this criterion was first used consistently by Tournefort. Although he admits that Linnaeus did much to increase our knowledge of the genera, Lamarck criticizes, quite correctly, the fact that so many of the Linnaean diagnoses appear to be based on a single species of the group. It is of importance in these diagnoses to admit nuances and to allow for exceptions by using words like 'ordinarily,' 'mostly' etc. Lamarck recognizes the philosophically different attitude of Linnaeus, whose genera were put forward as natural and objectively existing in nature. Lamarck'sreaction is that Linnaeus "a prononc' l'anatheme contre ceux qui assureroient que les genres ne sont point dans la nature...," and that it was much simpler for him [Linnaeus] to support his opinion by the so-called axioms and extremely laconic maxims with which he filled his Philosophia botanica and Critica botanica, than by solid proof which alone could convince those who were not impressed by mere authority. Lamarck puts his finger on Linnaeus's weak spot: He, and many others, maintained that nature makes no jumps "ce qui signifie, si je ne me trompe, que la serie de ses productions doit etre nuancee dans toute son etendue." This consideration alone, says Lamarck, makes it absolutely impossible to find well established, concrete lines of separation in nature between groups such as genera; he is therefore firmly convinced of the nominalist nature of all taxa above the rank of species. Lamarckis an equally firm believer, at this point, in the concept of a single series and in the principle of plenitude. Like Adanson, he has no doubt that the gaps in the series will be filled by future discoveries. Lamarck maintains that Linnaeus's dogma of the natural and objective character of genera pro410 TAXON This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions VOLUME 20 vided him, Linnaeus, with an excuse for the arbitrary nature of his generic circumscriptions.The natural character was invoked to authorize the adoption of many unsuitable assemblages of species. Linnaeus's axiom that the characters of a genus must be taken exclusively from the fructification is not accepted as an abolute rule by Lamarck: in certain natural families ("which are nothing but large uninterruptedparts of the series of plants") like the Umbelliferae or the Labiates, this axiom becomes sometimes unworkableand one will have to use other characters as well. Linnaeus's essentialist statement that the genus constitutes the character, not the character the genus (Phil. bot. no. 169) is firmly rejected. The genera are small parts of the great series and are characterised by form relationships; they remain a product of 'art' and not of 'nature.' The circumscriptionof genera must be highly pragmatic. The primary reason for establishing genera is to reduce the number of principal names to remember. a number which would be enormous if every species had a different single name. Botanical genera are like the constellations of stars: they are devices to facilitate classification. It is therefore necessary to make the genera neither too large nor too small. The creation of many monotypic genera is impractical, just as is the recognition of too-large genera such as Cavanilles' Geranium or Linnaeus's Lichen. Families (Enc. 2: 448) are again a device to help our imagination and to facilitate our insight into the form relationships. A single uninterruptedseries is difficult for the human mind to grasp; without interrupting the chain it is nevertheless necessary to recognize groups of genera. Lamarckdoes not believe that nature is subdividedlike an army into Brigades, Regiments, Bataillons, Squadrons, etc. (a clear reference to Linnaeus's Philosophia botanica); he emphasizes again that such divisions, however useful and necessary, are artificial. Lamarck accepts a gradual, almost continuous increase in organization, but no hierarchy. The scala naturae is to him not really a scale with steps, but a slope paved with species. The idea of a hierarchy objectively existing in nature is not necessarily the same as the recognition of differences between 'high' and 'low.' The concept of hierarchy in taxa is very old and can be traced back to two different sources. The first of these is the usefulness of hierarchical classification as a logical device, formalized for instance by Aristotle. The second source is man's age-old cosmic view that the world of beings reflects the situation in human society, where we have an hierarchical order based on differences in power. The Enlightenmentwas anti-authoritarianand it is understandablethat such prominent exponents of it as Buffon and Lamarckshould deny the objective existence of a power hierarchy whether in society or in nature. Rationalizing this, they shifted to the far end of the concept of plenitude: continuity. This shift was necessary before they could take the second step: continuity in space (here the world of living beings) reflects continuity in time. Spatial relations were always open to division and hierarchy; the only fundamental condition in human life which is not open to them is time. All living beings are equally subject to the final consequence of time: death. The egalitarian and antiauthoritarian tendencies in society are thus reflected in science. They were conditional for the recognition that a different degree of biological organization may have a historical basis. A historical - and therefore real - connection between 'high' and 'low' is not acceptable to the hierarchical mind. It is sufficient to point out the resistance against the idea of the descent of man to show how essential this breakdown AUGUST1971 411 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions Etr orga•que•svivans, afujcts a r NIMAUX.E.Tu.mimes. 1. LES QUADRUPEDES. 1. Terrearesenguicuks; & ui rprodi re LES POLYPETALEES.r; Thalamiflores. 4. Terreftres oguls. Caliciflores.2. Fre&iflores. , LES MOOPATAAESE ..T.X. E Y&ilres. x. C..i.i...res 2. Thalamifkre 3 A.Marms. LES OISEAUX. x. Terreftres. s. Aquatiques cuifes nues 1, Aquatiques nageants.- .. IES 4OMPOSAES . LES AMPRIBIES. * L Tetrapodes. S f t a facl Iiftindes.I Tubuleufes. a. Apodes. Ligulaires3. LES INCOMPLETTES... Thalamiflores. . .LES POI&ONt 1. Cartilagineui. ... 4. Caliciflores.2. 3. 2.EpineuxToDiclynes. Gynandres.4. LESINSECTE ILESV?UNILOBE...,.. Fru&fltres r. 1.'Tetraptires. u4.Diptires. rhalamidlorea;2., S. .. J.Aptbres. . L.E. . V Nuds. 4. Teftacs. ;. 2otanique.Tomel, FIG. 3. 3. Lithophytes;i.> 4. Zoophytes. LES CRYPTOGAMES . Epiphyllofpermes.i. Urnighres.2-,i 3. embraneues% Fongueues.4. Lamarck's botanical and zoological series as presented in the article Classes of the Encyclopddie methodique, Botanique 2: 33. 1786. of social class structure was in making scientists shed their essentialist and scholastic past. In this early botanical work Lamarck admits only a single unbranched series of plant taxa, parallel to one for the animals. In later years, after he had understood the historical background of this image of a chain, Lamarck could also admit the possibility of further branching and of the occurrence of dead-end series in the course of evolution. In his article on classes in the Encyclopddie (2: 29) he develops his thesis of the two parallel series in nature. Even though his concept of two series of animals and plants is still timeless and only a descriptive picture of the order of complexity of organization, it is of the greatest importance because it was a decisive step in the breakdown of the old chain of being. All through the ages the 'chain' had been a single scale from mineral via vegetable, animal and human to less tangible metaphysical entities. Lamarck's important statement on species, quoted above, showed his awareness of a difference in principle between the inorganic world and that of living beings: the birth of biology. Now, with the establishment of two parallel chains of living beings, the old 'chain' is broken a second time. Lamarck describes the existence of a third chain in this article on classes in the Encyclopedie. This third chain is that of the inorganic world. All non living substances on the crust of the earth are said to be of organic origin. In time they degenerate into simpler components: at the beginning stand the highly organized organic remains of dead animals and plants, at the end substances that strike a light on flint, such as quartz. 412 TAXONVOLUME 20 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions Lamarck's idea of organic evolution emanated from his views on the formation of the various mineral substances of the crust of the earth. The key to the development of Lamarck's evolutionary thinking lies in his two tables in the article Classes of the Encyclopedie (2: 33, 34; see figures 3 and 4). The first table, of "organic living beings, subject to death, and which have the faculty to reproduce themselves" shows an obviously intentional close parallel between the animal and plant series. Both have six classes, arranged in order of decreasing complexity of organization. This arrangement alone is sufficient proof of the absence of the time element in this representation of the marche de la nature. Each class is subdivided into 2-4 sections which, for the plants, have much in common with the groups distinguished by Tournefort. It is interesting to note that the polypetalous plants are considered to be the most complex and that the Unilobees (monocotyledons) are placed closest to the Cryptogams. It should also be noted that Lamarck presents a zoological classification with several original elements, years before his appointment to the zoological chair at the Museum. The botanical sections comprise the 94 families which are mainly in accordance with those published later by AntoineLaurent de Jussieu. Lamarck remarks that it is preferable to go from the complex to the simple rather than the other way around, because the information on the more complex organisms is more complete. On the other hand he is fully aware that "we are still far away from a complete knowledge of the real but does not specify order of gradation of which we realize the importance... ," that importance. His classes and families are typically 'natural,' based upon a great variety of characteristics; the sections, as later with Jussieu, are much more pragmatically based on relatively few characters. The main guideline of the classification is, however, "the order of gradation with respect to the perfection of their organs." The whole scheme, with its forced symmetry, shows an emphasis on form rather than contents, a typical feature of 'classical' neo-platonic thinking. The ladder may be a double one, but it remains a static thing. The main criteria for the botanical system are derived from 'essential' characters(reproduction) and the division of the cryptogamsis extremelyprimitive. The really advanced element of the system lies in the arrangementof the families, which is based on a great variety of criteria. Lamarckknows that this arrangement of the families is not yet ideal; with respect to the main classes, however, he says that the order "ne nous paroit nullement susceptible d'un changementqui puisse etre plus convenable." In sharp contrast with these comparative morphological series is the scheme of the "inorganic beings, without life, and produced by the successive alterations of the composite substances which have been part of living beings." (Fig. 4). This table illustrates the genetic relationships among mineral substances. Lamarck stresses that it represents an entirely new concept; "that all minerals are the true products of successive alterations to which the debris of organic beings are subjected; that these minerals are not at all the result of direct formation, nor of successive reproduction, but that on the contrary they are the result of the continuous alterations to which the debris of living beings are sujected, alterations which transform them successively in as many diverse components as there are minerals known." "One sees from this table the infinite distance between living beings and inorganic substances, and the need not to present beings of such different character in a single continuous chain." The fundamental difference between minerals and living beings is not only that the latter are maintained by successive reproduction, but also that minerals are formed indirectly and from the 413 AUGUST1971 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions IErl s inorganiqjuesfans vie, & produitspar lks altdraeionsf'cflives des ftances compofiesqui oncfairpartiddes isres vivans. :Terreauanimal Terreauvegita v gtaat Terre4uatimal 7Terreau es Crsiflacls, &e. desCime.&des Voiries; , des1Marais. des ChamrpsdesBois Teac Qcoquilliere.. Cra" Tourbe. 4arnes. Bitumeso. .Sonfres? `vi:PcrrcS ~ca~c; " Mgrbres, . Nitre. -l Borax. Gyp?es. Agl> AIUn. Abte.Alkls taIre$, Shits. Tals it Pyrtes. SSpata ( Terre franche. SMaths Pieere Schorbs. rneutiere. CrIftaux Cailloux" A gemmes, PierresA ath.. Feld- de Rohch. ju.tnOWit FIG. 4. Lamarck's scheme of the inorganic substances as produced by successive changes from substances of organic origin. remains of living beings. The mineral chain is one in time, the plant and animal chains are - for the time being - arrangements in space only. The mineral chain goes away from life, while the others represent simply an increasing complexity within it. This contrast of physical and organic nature is dialectic and indicative of a new philosophical trend. Only life can synthesize; inorganic nature destroys complexity. Implicit in this view, which we find all through Lamarck's work, is an interesting - although primitively expressed trend of thought in which life is a phenomenon maintaining organization, a high energy level, away from chaos. In modern terminology: life maintains a minimum of entropy; inorganic nature tends to reach maximum entropy. Needless to say that Lamarck does not use these words; they are but a twentieth century rendering of what were with him embryonic but strangely lucid ideas emanating from a primarily deductive, and in details incorrect, cosmology. The thesis of the organic origin of all minerals is obviously untenable: to state that metallic ore, granite and basalt are derived from organic substances is too far fetched. Lamarck was a geologist whose field experience was limited essentially to the chalk and tertiary beds of the Paris Basin (Carozzi, 1964). 414 TAXONVOLUME20 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions The sweeping generality of the thesis detaches it from reality; the importance of it for Lamarck's genetic thinking, however, is capital. His ideas on peat and coal formation, even though not completely correct, are modern for the period and found support in, for instance, Bruguieres' Choix de mimoires d'histoire naturelle of 1792. Lamarck'sideas on this subject were a first sketch of what would later become a main theme in his Hydrogeologie (1801) in which he presented an actualist and dynamic view of the history of the crust of the earth. Here, in the Encyclopedie (1786), we find a first proof of Lamarck's deductive reasoning and leaning towards a dynamic view of nature. The classification of living beings, with its two almost symmetrical 'scales,' may still be essentially static; the whole article on classes promises a new approach. The 'biological' emphasis is amazing: until then no one had given such a central place to the production of organic material in any cosmology. Lamarckevidently enjoyed considerable freedom in the choice of his material for publication in the botanical part of the Encyclopedie. He had formulated his thoughts on the overall organic origin of all mineral substances for the first time in a manuscript dating from the years 1776-1781, published later (1794) under the title Recherchessur les causes des principaux faits physiques. In this work he relates (2: 366) that he had not been allowed to publish these ideas as he wanted in the first volume of his Flore franqoise, but that he was able to put them into the Encyclopedie. The argumentation in the Recherches is much more elaborate and touches also upon the origin of living beings in general. His conclusion with respect to this fundamental question is still negative: "Or, je crois qu'il est aussi impossible a l'homme de connoitre la cause physique du premier individu de chaque espece, que d'assigner aussi physiquement la cause de l'existence de la matiere ou de l'univers entier." Life is something sui generis but Lamarcktakes an importantstep by clearly stating that he ignores the origin of life in general as well as that of individual species: he moves away from a creationist attitude. The species concept, however, is still static. In his article Froment (wheat) in the Encyclopedie (2: 557) Lamarck explicitly states that the essential charactersof a plant are not known to change in cultivation. There is so far no example that as long as the reproductiveorgans remained functional they have taken on characters which were not those belonging to the plant to which they originally belonged. This point of view is the same as that of Adanson (1772) who, after first believing that such change was possible, later reverted to the constancy of species. Lamarck'sunusuallykeen insightinto other generalbiologicalmattersis exemplified in the Encyclopedie, for instance in his article Foret. He discusses the need for a controlled policy of conservation of forests and adds that the destruction of the woods and forests for the benefit of special, annually highly productive crops, leads to a gradual loss of fertility. In the end the land will become barren, human habitation will be abandoned,andall otherformsof life will ultimatelydisappearas well. Lamarck's analysis of the r6le of forested areas with respect to prevention of erosion, desiccation and loss of arable soil is modern. The forests are correctly seen as the great producers of biomass with a decisively temporizing influence on the climate; they are indispensable to maintain that important source of wealth and well-being any country: its good arablesoil. Lamarck'sEncyclopediecontains a wealth of material and ideas. We are again and again struck by the consistency of his cosmology, or rather of this theory of the earth. In this theory there are remnants of age-old patterns of thought, but there are also indications of new trends. Some of these ideas were further 415 AUGUST1971 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions elaborated during the tumultuous years that followed Lamarck's appointment as a zoologist. At first there were few outward signs of his manifold activities, but the new century brought proof of an almost explosive outburst of creativity in the form of a great number of publications in each of Lamarck's spheres of interest. From gradation to descent (1800-1803) The publications by Lamarck of the years 1800-1803 display an astonishing variety. First there are several meteorological publications including the Annuaires meteorologiques, then there are the Systeme des animaux sans vertebres (1801), the Hydrogeologie (1801), the Recherches sur l'organisation des corps vivans (1802), the first instalments of the long series of the Memoires sur les Jossiles des environs de Paris, the Histoire naturelle des vig~taux and the various opening lectures of the zoology courses at the Museum. The last years of the century must have been filled with feverish scientific activity in all these fields. It must have been during those years that Lamarck finally abandoned his belief in the fixity of species and came to the formulation of his transformist views (see on this also Gillispie 1956). His continuous interest in fossils, his work on the influence of water on the surface of the earth, his attempts to understand geological time, and his realization of the repeated changes of environmental conditions in geological time at specific places must all have contributed to the further evolution of his thoughts on the dynamism of nature. In his views on the genesis of minerals, contrasted with the series of animals and plants. Lamarck had come very close to grasping the significance of natural classification. The main factor responsible for his shift from 'gradation' to 'descent' in living organisms parallel to degradation in time of organic compounds to simple minerals must have been his keen insight into the importance of the time factor and, to a lesser degree, of environmental conditions. Landrieu (1909) has tried to follow this development step by step in Lamarck's publications. The first really clear statement of a theory of organic descent is found in Lamarck's Discours d'ouverture du Cours de l'An VIII which was delivered on 11 May 1800 (reprinted in Lamarck's Systeme of 1801). This lecture is the birth certificate of transformism. The first - very long - sentence contains the statement that Lamarck will explain in rough outline the order which nature seems to have followed in forming its animals. He further states that evertebrate animals, more than any other living beings, are of interest to the philosophernaturalist because they confront him with the simplest organic beings, "those perhaps by means of which nature started and from which, with much time and favorable circumstances, she has formed all others." Time has been at the disposal of nature without limitation, the circumstances arise from the influence of the climates, ecological diversity, etc. As a result of these diverse influences the 'faculties' change and fortify themselves through use. The changes are conserved imperceptibly and, in the long run, propagated by generation. The essence of Lamarck's theory of descent is contained in this early lecture; he even mentions the classical example of the feet of the swimming birds. The debatable mechanism of descent (acquisition of characters by individuals or by populations?), now usually considered the core of Lamarckism is not really so important. Very often excellent theories have been formulated on the basis of faulty proofs, inadequate hypotheses, or unclear semantics. The history of science is full of such cases. Lamarck's main thesis is that organisms (not necessarily individuals or the immediate offspring of specific individuals; LaTAXONVOLUME20 416 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions marck had no word for the concept of population), change with time, and consequently species are not constant.* In this same year, 1800, Lacepede published a Discours sur la durde des especes as an introduction to the second volume of his Histoire naturelle des Poissons in which he states that the species is only an abstraction, 'a collective idea,' and in which he analyses the importance of extinct species. He comes to the conclusion that "a species can undergo so many modifications ... that it will ultimately be more distant from its original state than another species: the species has then undergone a metamorphosisinto a new species.. ." A complete knowledge of the past will reveal "the successive modifications of organized matter ..., the past will serve us to complete the idea of the present." Lac6pede did not further elaborate these essentially Buffonian ideas. This was left to Lamarck. The reason why both Lamarck and Lace3pde came forward with their ideas more or less at the same time in the year 1800, and not in the earlier years when these ideas must have grown in their minds, may perhaps be found in the dominating influence of the old director of the Museum, Daubenton, who was evidently much opposed to this type of thinking. Daubenton died on the first of January 1800. Lamarck, in his famous lecture, points out that the series of animals and plants reflect the increasing complexity of organization. With the animals the starting point may be "la monade qui, pour ainsi dire, n'est qu'un point arrive'e. . . " in the vegetable kingdom it is the "byssus pulverulens"(lichenized algae, now often called Lepraria) or "la simple moisissure," (mucor viridescens, according to Lamarck himself), which seems to be the minimum of vegetable organization. Almost equally important is Lamarck's change of the picture of the animal and plant series. As long as the species were considered to be constant, he had adhered to a linear series of species, an extreme view of the scala naturae. Now it is necessary to modify this representation:there may be species brought about by special circumstanceswhich stand a little aside from the regular flow of evolution and which are side-branchesof the main stem. "By that gradation nuancee in the process of increasing complexity of organisation I do not mean to speak of a linear series with regular intervals of species and genera: such a series does not exist, but I speak of an almost uniformly gradated series of masses principales such as the big families, a series which certainly exists in the animal as well as in the vegetable world, but which as far as genera and especially species are concerned, forms many lateral ramifications of which the ends constitute really isolated points" [translation]. Lamarck has now come very close to the modern picture of a stream of organisms, linked by their changing genomes with branching dead-ends, but also with far-reaching continuity. He leaves no doubt that the natural system is a reflection of this biological process. At last the great social ideal of the Enlightenment, progress, is reflected in biological science; at last, also, we witness the complete breakdownof the timeless chain of being. The main elements responsible for this development of Lamarck's thoughts are his early work on the organic genesis of certain deposits (unduly extrapolated into an overall theory of mineral descent) and his more general preoccupation * Bernard Germain French Etienne de la Ville, comte de Lac6pede, 1756-1825, zoologist, proteg6 of Buffon, active revolutionary, president of the Legislative, Professor of zoology at the Museum d'Histoire naturelle, ennobled by Napoleon and by Louis XVIII. AUGUST1971 417 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions with the history of the crust of the earth, including the interpretationof fossil remains. This development is illustrated in his Hydrogeologie of 1801 better than in any of his earlier publications. This extremely rare book (Lamarck published it at his own expense and sold very few copies) is now available again in an excellently annotated English translation (Carozzi 1964). The relatively short text is eminently readable and shows all facets of Lamarck's thinking: often wildly deductive and spontaneously generalizing 'a la Buffon, but more often surprisingly modern in the tentative explanation of many geological and biological processes. Lamarck shows himself to be a consistent geological uniformitarian (actualist) by explaining the past in terms of the present: "this is the path of Nature resulting from her well-establishedprocesses and finally demonstrated by the phenomena continuously occurring before our eyes." This geological uniformitarianism is also biological. In order to explain the world of living beings one can restrict oneself to "phenomena continuously occurring before one's eyes." Lamarck had already tacitly condemned creationism in his earliest work (published 1794) by stating that the precise details of the origin of species might remain obscure and by clearly not invoking metaphysical interventions. A similar pattern was found in the 'three series' of the Encyclopedie: there was no room for beings beyond man. The biological uniformitarianismof the Hydrogeologie is the basis of Lamarck's transformism. It had been foreshadowed halfheartedly by such biologists as Buffon and Adanson but these scientists had had to pay lip service to creationism. The realizationof the immensity of geological time had also come as a result of Lamarck'swork on fossils and on general geology. He had a clear understanding - certainly in part because of Buffon's earlier work in this respect - of the climatic significance of fossils, for instance as shown by the discovery in temperate regions of animal fossils indicative of tropical environment. "Oh how very ancient the earth is! And how ridiculously small the ideas of those who consider the earth's age to be 6000 odd years!" . . . "The great age of the earth will appear greater to man when he understands the origin of living organisms and the reasons for the gradual development and improvement of their organization. This antiquity will appear even greater when he realizes the length of time and the particular conditions which were necessary to bring all living species into existence. This is particularlytrue since man is the latest result and present climax of this development, the ultimate limit of which, if it is ever reached, cannot be known." ... "Time is insignificant and never a difficulty for Nature. It is always at her disposal and represents an unlimited power with which she accomplishes her greatest and smallest tasks." [translations Carozzi.] We find similar thoughts with Lamarck's colleague and friend Lacepede, whose Histoire naturelle des poissons (1800) was mentioned above. Lacepede also stresses the immensity of biological and geological time: ". .. those pyramids which we call antique because we ignore how many millions of generations disappeared before the moment when their height became a rival to that of the mountains. . ." Lac6pede defined nature also as 'space, time and matter.' At this point in the developmentof biological ideas we cannot ask for more. Man's place with respect to geological time and his descent from non-human beings is clearly stated. No wonder Lamarck and Lac6pde found no echo for their theory. Lamarck especially had exposed himself somewhat too conspicuously with his theories of chemistry and physics which had given him the 418 TAXON This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions VOLUME 20 reputation of a wild speculator. The Hydrogeologie remained almost undistributed; practically the whole impression was found among his possessions after his death. Lamarck's ideas became more widely distributed through various publications in the Annals of the Museum and, in part only, through his Philosophie zoologique. The idea of the vast antiquity of the earth and the implications of this antiquity for geology and biology remained unaccepted for a long time. In the first part of his Memoires sur les Jossiles des environs de Paris (1802), Lamarck discussed again the importance of the fossils as indicators of slow but continuous change in climate with respect to every point on the earth's surface (see also Carozzi 1964, p. 75). Man has a natural tendency to disbelieve all the facts presented by the fossils. He is like insects with a life span of one year living in a building and trying to describe its history. "For twenty-five generations there has been no change: hence the building is considered eternal ..." The fundamentals of the Hydrogeologie were written for the first time, says Lamarckhimself (p. 173), on 21 Pluviose of the year 7 (9 February 1799), which shows that this intriguing book reflects Lamarck's state of mind during the crucial years between 1797 and 1800 when he came to a major change in his geological and biological views. Lamarck himself was under no illusions as to how his work would be received. The last words of the book are: " ... if respect for the idol that should be toppled prevents one's eyes from opening, I will say: Let those who prefer to be deceived be thoroughly deceived." Lamarck's switch towards transformism had one tangible result in botany. His last botanical publication, almost universally overlooked, is the general introduction to the botanical part of a new set of Suites a Buffon, the eightyvolume (sic) Cours complet d'Histoire naturelle for which Lamarck and Mirbel wrote the Histoire naturelle des vigitaux (15 volumes, Paris 1803; reissued 1825). Lamarck wrote the text of the first two volumes: an introduction containing a brief history of botany, terminology and physiology, and a discussion of the general principles of classification. The greater part of these two volumes is an almost word by word copy of corresponding entries in the Encyclopedie, published 15-20 years earlier. Evidently Lamarck spent little time on the compilation of his final botanical publication; there are, however, a few places in which the text is updated such as for instance in the chapter on the "natural and methodical distribution of plants" which contains a discussion of the natural system. The text is familiar, with its emphasis on the marche de la nature but the text on the linear arrangementis in accordancewith that of the famous Discours d'ouverture of 1800. The natural order is the "distribution conforme a la marche de la nature et au plan qu'elle a suivi en produisant les &tresqui les composent." [p. 261] The system proposed still has much in common with that of the Encyclopedie, but it differs fundamentally because the order is reversed. The cryptogams (divided into two classes, 'Aphyll6es' and 'Cryptospermes') now stand at the beginning rather than at the end: the phylogenetic system has made its appearance. It is one thing to propose a theory of descent and to formulate the conviction that the natural system is a reflection of phylogenetic reality - a typically 'systematic'statement-; it is anotherto formulatea theory about the mechanism of evolution and the origin of life. Obviously, the first is a condition for the second; the mental process of accepting 'descent' or evolution is fundamental. The explanation of how it all same about is only in part the subject of systematics. Lamarck's theory of hereditary adaptations to changing environmental conditions need therefore not be discussed in great detail; this has been done AUGUST 1971 419 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions at great length by others. However, for the development of ideas in systematics the process of admitting that living beings have a history and that the present picture has to be seen against this historical background is of overriding importance. For nearly twenty centuries the ro1e of time had been neglected. The publication of this new classification in 1803 is epoch-making because it is the first major classification of the plant kingdom based upon a theory of descent. Obviously the classification itself is purely morphological; the order is determined by an idea, the factual contents by morphological experience. This is a far-reaching consequence of Lamarck's actualism. In geology, as in biology, Lamarck held that historical development can be explained without invoking other causes than those we can witness today; he is therefore logically correct when he advances a comparative morphological series as an image of historical development. This theoretical basis of classification is further worked out in the Philosophie zoologique. It would be wrong, however, to think that Lamarck's ideas had any impact on biological thought in his time: they had none. Biological philosophy and classification (1803-1809) The account of Lamarck's ideas in botanical systematics could end at this point: the publication (1803) of his last overall classification of the plant world. ' Everything had been said: actualism' had been accepted as a main theoretical basis, 'descent' had been made the theoretical - though not factual - basis of classification, a mechanism of evolution had been suggested, the fixity of species had been abandoned. However, the development of these ideas, the refinement and precision of their arguments and the way the ideas were received by Lamarck's contemporaries are also part of the history of ideas in systematics. The emphasis was almost exclusively on zoology, but there was still much to say on general systematics. Lamarck's main publications after 1803 deal with fossils, evertebrates and theoretical biology. There are 33 Memoires sur les fossiles des environs de Paris (1802-1806). Numerous brief descriptive notes on new evertebrates, as well as the great Histoire naturelle des animaux sans vertebres (7 volumes, 1815-1822) are proof of Lamarck's continuing output as a descriptive biologist. The Philosophie zoologique (1809), the Introduction (1815) to the Histoire naturelle and the Systeme analytique des connaissances positives de l'homme (1820) are his major theoretical works of late middle age. In addition there are a number of interesting theoretical articles in the second edition of Deterville's Nouveau Dictionnaire d'Histoire naturelle (1816-1819). The Philosophie zoologique could also have been called Philosophie biolo. gique, because many of the considerations are of general biological importance; references to botany are not at all rare. Lamarck's primary assignment, however, was zoology, and many of his most important ideas stem from his zoological work. Before discussing some aspects of this well written and stimulating book, however, it should be stressed that one does Lamarck an injustice by judging him from this book - his best known - alone. His ideas on the dynamism of nature and on the possibility to understand the past in terms of the present developed as a result of all his activities in the natural sciences, and not least from his geological work. The fundamental change in Lamarck's thinking over a period of years around the turn of the century is the gradual realization of the important r8le of time in all of nature's processes. His early, keen insight into the absolute length of time involved in geological processes stands at the beginning of his biological philosophy. Many of the thoughts expressed in the Philosophie zoologique can be found in a more elementary form in his earlier 420 TAXON VOLUME This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions 20 publications, especially in his opening addresses of the zoological lectures at the Museum (Lamarck 1800-1806). The Philosophie and the Introduction (1815) contain the synthesis of Lamarck's biological thought and express his ideas most eloquently and precisely. To regard Lamarck'shypotheses on the sheer mechanism of evolution as the main feature of his thought would be an injustice to his much more important general theses. At this phase in the history of biology the science had simply not yet developed far enough to allow for a more sophisticated approach, such as, for instance, that of Darwin and Wallace. Experience and experiment fell short of the needs for an inclusive theory. The main themes of Lamarck's biological theory had to be deduced from the factual knowledge available to him. It is therefore more important that he realized the existence of phenomena such as adaptation, variation, speciation and evolution than that his picture of the mechanism was, on the whole, untenable (at least if restricted to individuals). Lamarck, like Buffon, was a deductive genius with great factual knowledge but without the urge to test the deductions by means of experiment or to widen his horizons through travelling. His deductions, however, are proof of a powerful scientific imagination, able to produce trains of thought that would stimulate more refined research by others. The deductions also broke through the boundaries of much that was still sacrosanct, and they thus played their r81e in freeing the human mind from an unscientific bondage with the past. Lamarck never used the word 'evolution,' and even the word 'descent' does not figure to any extent in his books. 'Descent' would really not have expressed his main thesis: if he had wanted a word like that he would rather have used 'ascent.' Instead, his terms are 'complicating organization' and 'successive production.' When Lamarck speaks of 'distribution' he refers to what is now usually called taxonomy; his 'classification' refers to the delimitation of taxa, not to their mutual arrangement (which is 'distribution'). The words 'histoire naturelle' in the title of his main work on evertebrates comes closest to our 'systematics.'The main criterion for the 'distribution'(taxonomy) is the 'marche de la nature' or the 'echelle' (ladder of nature), originally identical with the platonic concept of a static smooth gradation of form relationships, but later an expression of the actual course of events in the history of living beings. At first the ladder was a flight of minute steps (species), ultimately it became an escalator, moving all the time, not in an endless chain but picking up newly created living matter at the bottom. Lamarck's main thesis on taxonomy is that the natural system (based on the greatest possible correlation of morphological characteristics) is identical with this marche de la nature: this holds for his early - timeless - thinking as well as for his later - evolutionary approach.The logical need for this identity was disputed by many, and Lamarck tried to prove it by pointing out the actual course of events. This 'proof' was, of course, not empirical but deductive.This identity, however, was, for Lamarck, an almost undisputable fact. In his later years his descriptive morphological work on evertebrates made him admit the possibility of a marche which was not a ladder, but a mainstream with sidestreams. The word embranchement is later introduced to account for this phenomenon of evolutionary divergence. Finally, the word 'nature,'"so often pronouncedas if it dealt with a particular being," can, for Lamarck,be only the totality of its constituents: "1. all existing tangible objects, 2. the general and special laws governing the changes and situations [sic] to which these objects can be subjected, 3. the movement, generally and diversely distributed among the objects, perpetually kept up or AUGUST 1971 421 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions reborn from their source, infinitely varied in its products and from which results the admirable order of things presented to us by this whole." Here again we find a new element in the definition of an age-old concept. Nature had generally been defined solely by items 1 and 2. The third, rather involved component [here almost literally translated, Phil. Zool. 1: 359-360] is Lamarck's profession of faith as a scientist. Nature is dynamic, evolving, creative, infinitely diversified and still in the process of diversification. Strictly speaking, item 3 simply identifies one of the major laws implied in item 2; in fact, however, its specification embodies a new view of nature: the discovery of the intrinsic value of the dynamics of diversity. This is the final outcome of the principle of plenitude, the definitive shift from an immutable, self-sufficient uniformitarian universe to a world characterized by creativeness, by Kant's temperal ego, by Schiller's Stofftrieb, by the realization that an individual is part of a stream of ever-changingforms of life. Lamarck, in short, pits scientific romanticism to scientific classicism. The first elements of nineteenthcentury positivism are contained in this significant definition of nature. Lamarck describes the development of his thougths in the avertissement to the Philosophie zoologique. A survey of the precepts and principles relating to the study of animals would be useful, says Lamarck, also to other natural sciences. While recopying and describing the most general rules applicable to the observed phenomena, Lamarck felt himself gradually formulating considerations of the greatest interest to science as a whole: "How, really could I envisage the singular degradation which is found in the complexity of animals, going from the most perfect to the most imperfect, without trying to determine how to explain such a positive and remarkable fact, for which there are so many proofs? Had I not to think that nature had produced the various living organisms successively, going from the simplest to the most complex, because when we retrace the steps of the animal ladder the organization of the animals is seen to become gradually more complicated in an extremely remarkable * way?" This thought was stimulated by the realization that simpler organisms had fewer or no specialized organs, and that further complexity meant further morphological specialization and division of functions. The origin of life and of the process of increasing complexity were now seen in close connection with the diversification of organs, the adaptation of living beings to their environment, and to changes in their environment. The book deals with much more than the ideas of direct interest to us here: many biological, especially physiological, phenomena are dealt with, including subjects such as human intelligence and morale. These treatises were sometimes far reaching. A statement, for instance, that the body and the mind have undoubtedly the same origin was not exactly in agreementwith contemporaryreligion and philosophy. A discussion of this part of the book, however, would be less relevant to our purposes, even though Lamarck'swhole philosophy supports his revolutionary ideas on biological phenomena such as adaptation and evolution. The word 'degradation,'mentioned above, is rarely used by Lamarck.Buffon used it partly as a subterfuge; in Lamarck'stime this was no longer necessary. His thoughts might not be acceptable, but the church could no longer press him to publish a refutation of his scientific postulates. There had been, after all, a French revolution, and Lamarck is in many respects a typical product * This and the following quotations are actually rather free translations of Lamarck's original, not based on any of the existing translations, aiming at a rendering of Lamarck's ideas in our contemporary idiom and terminology. 422 TAXONVOLUME20 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions of this Revolution and not just of the Enlightenment.His Philosophie zoologique, however, appeared in a period of restoration of much that the Revolution had tried to abolish, and he himself had come under suspicion of remaining too revolutionary. When presenting his Philosophie to Napoleon, Lamarck received in return a snub about his meteorological ideas, which went against those of the scientific establishment of the Empire. The word 'progression,' with all its overtones of the Enlightenment and early romanticism, is much dearer to Lamarckthan degradation. Life was a process of slow progress in organization towards perfection: "with respect to the bodies that enjoy life, nature has done " everything little by little and successively . . . Lamarck announces as a new edition of which The Philosophie zoologique, his previous Recherchessur l'organisation des corps vivants of 1802, is divided into three parts: the first dealing with systematics in the modern sense, the second with the origin of life and the nature of the life processes, the third with the origin and nature of 'sentiment'and intelligence. Our analysis is of the first part only (chaptersi-viii). Lamarckian systematics in the Philosophie zoologique (1809) Lamarck's definition of systematics (called histoire naturelle) is contained in the title of the first part of the Philosophie: "considerationson the systematics of animals, their characters, relationships, organisation, taxonomy, classification, and species." The term histoire naturelle is as protean as its counterpart nature; every phase of western culture has had a definition and circumscription of its own. It is unusual, though, to find it defined as closeley to our present concept of systematics as is the case with Lamarck. Lamarck uses also another term in a somewhat unorthodox manner: he speaks of the 'part of the art' of the natural sciences, where we should now simply use the words methods or methodology. It is very important, says Lamarck, to distinguish between the art and the laws of natural history. Another distinction lies between economic and pure ('philosophical') biology. The natural sciences, and biology in particular, found their origin in economic need and practice. 'Pure' followed 'applied' science. In the eighteenth century the new element of pleasure (agrement) was added as a justification for studying 'natural history.' 'Pleasure' in studying botany was of educational importance. The Enlightenment strongly promoted the utilitarian and didactic features of the natural sciences in order to spread knowledge at a popular level. Pleasure and usefulness may seem contradictory aims; to the philosophers of the Enlightenment, however, incipient revolutionaries as they were, these aims should be combined in order to improve or change the structure of society. This change could not come from the poor and illiterate, but had to be effected by the creation of a non-elite intellectual middle class. Botany played an important educational ro1e because it was pleasant and appealed to the reverence for 'nature.' It is interesting to see that more than a century later the idealistic socialist type youth movements would emulate this eighteenth century ideal for almost the same reasons.* Systematic botany, in a primitive ro1e of innocence and beauty, thus played a modest part in improving la condition humaine. Lamarck himself demonstrated this attitude in his article on botanique in *) A conspicuous example of such a group was the Netherlands youth movement for the study of nature (N.J.N.) which was especially active between the two world wars. Many of today's Dutch biologists belonged to this group in their early years. AUGUST 1971 423 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions the first volume of the Encyclopedie. Among the main merits of botany, that of 'object of pleasure' is given a conspicuous place: "how many powerful motives are united [from the point of view of agrement] to invite us to study a science so capable of giving us such a pure satisfaction, so worthy of an honest soul ..." The eulogy continues, with many words. How different is the attitude of 1809. Now the pleasure is just mentioned incidentally, and as an historical cause, but "the purely scientific interest makes us eager to know nature itself in all its products, in order to grasp its development, its laws, its mechanism, and to enable us to form an idea about everything in existence; briefly, this scientific endeavour procures that type of knowledge to which the true naturalist aspires." Biology is no longer the special subject of leisurely study by children and ladies, leaving the firm, but more directly useful branches of science such as theology, law and [military] technology to man. With the romantic movement in science biology comes of age. The main features of biological methodology or 'art' - as opposed to the 'true laws of nature' - are systematic classification, the higher taxa and nomenclature. Here Lamarck sticks to his eighteenth-centurynominalist guns: there is nothing natural about classes, families and orders, the so-called systems are artificial conventions, and nomenclatureis - of course - only the handmaiden of science. None of these are found in nature: "... .among its products, nature has formed neither classes, nor orders, families, genera, nor constant species [sic], but only individuals which succeed one another and which resemble those from which they came. These individuals belong to infinitely diverse races [i.e. species, sic] which differ minutely in all forms and degrees of organisation but which remain constant as long as no causal agent effects a change." This is a modification of Lamarck's previous view on the nature of species: since species change in time, they are not 'real,' hence the extension of the nominalist attitude also to species. Lamarck had now reached the same stage as Buffon half a century earlier; for the latter - also for purely deductive reasons - the only real entities in life were the individuals. Each taxonomy which is not in conformity with the order of nature is artificial; "we are now perfectly justified in recognizing only a single order established by nature ... this order is the one in which all living bodies have been formed from the beginning." "This order is unique, essentially without division in each organic kingdom, and can be known by us by means of the special and general form-relationshipswhich exist between the objects which are part of those kingdoms ..." This unique order "will have to replace ultimately those systematic or artificial taxonomies which we have been forced to create in order to arrange the observed natural objects in a commodious way." Systimatique means to Lamarck, as it did to Adanson, in accordance with an artificial system. It has therefore an entirely different meaning from our present word 'systematics.' "In botany the natural method is extremely difficult to establish," because of the obscurity of the form relationship. A. L. Jussieu had made some progress and many families had been established; however, their overall taxonomic arrangementwas and still is a matter of dispute. "Truly, one has only found the beginning of this order; the middle, however, and especially the end, are still at the mercy of arbitrariness." Lamarck explains at great length that all classes are artificial. In zoology it might be difficult to believe that groups like mammals or birds are not natural; yet, this is actually an illusion caused by our limited knowledge of the animals that exist or have existed. The more men's knowledge increases, 424 TAXON VOLUME 20 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions for instance because of the exploration of regions like Australia, the less defined the divisions between the classes become. Sometimes the diversity of objects belonging to one class is so great that numerous subdivisions are needed at various ranks. However, one should be careful not to increase the ranks of the taxonomic hierarchy, and especially not to burden science with new names for all those intermediate taxa. "It is necessary to put an end to this abuse of nomenclature; without this, nomenclaturewill become a subject more difficult to know than the objects which one has to consider." If all organisms were known, any division would prove to be artificial. Looking at this statement against a historical background, and in the light of Lamarck'spreoccupationwith fossils, it is clear that his picture of the living world in space and time is one of a continuous stream in which the term 'masses' stands more or less for the genome. However, Lamarck emphasizes that this perfection of knowledge is far in the future. For the moment our classifications and delimitations of taxa are facilitated by the gaps in man's knowledge. "When will the naturalists agree to classifying their species in a uniform way?" They are alas seduced by the conviction that 'their' genera, families, orders and classes really exist in nature, and create their taxonomy on the basis of their opinions rather than on fixed pragmatic principles. Lamarck therefore never objected to the use of the Linnaean artificial system in his botanical works even though he knew that the natural system held more promise for science. It cannot have been difficult for him to follow the wishes of Pancoucke that the Linnaean system be used in the Illustration des genres. Nomenclatureis also part of the 'art,' and a much abused one. It is necessary, says Lamarck, to realize that many name changes are the result of taxonomic considerations, such as differences in the delimitation of genera, which are really quite superfluous. As a result of this constantly increasing number of taxonomic changes the synonymy becomes a heavy burden which one day may obscure the scientific considerations themselves. The result will be "... an immense maze before which one trembles, with reason, on entering." Characteristics (chapter II). Lamarck devotes a whole chapter to the "importance of the considerationof the rapports."The word rapport,as usual at that time, stands for formal - not genetic - relationship. "For living objects, one uses the term rapport between two objects that are being compared, for the features of analogy or resemblance, taken from the whole or generally from their parts but attaching more value to the more essential ones." The word clearly stands for 'form-relationship'and has none of the 'genetic' overtones of the English 'relation.' The natural form-relationships are for Lamarck the guide to the natural 'method,'to the ranks that are assigned to the main 'masses' that are encountered in taxonomy. They force the taxonomist to adopt the order followed by nature in bringing its objects into existence. The characteristics or features upon which the form-relationshipsare based are sometimes so pronounced that it is sometimes possible to identify a species on the basis of only part of the individual. This is of importance in fossil studies. However, there is a limit here, because the species change in time, albeit exceedingly slow, so slow that they escape the sharpest observer. As a result most observers conclude that the differences which they see have always existed. Some characteristics are more important than others; their importance is directly related to the degree of generality of their relevance. For animals the 425 AUGUST1971 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions characteristics derived from the interior organization are most important, in plants those taken from the fructification. This weighting of characteristics is arbitrary, and Lamarck doet not specify the reasons for his decisions, apart from his reference to the 'degree of generality.' In plants he suggests the following order of importance: 1. the embryo,includingcotyledons,perisperm,and the seed which containsthe embryo 2. the sexualpartsof the flowers 3. the non-sexualpartsof the flowers 4. the pericarp 5. the reproductive organswhichdo not requirefertilization. Lamarckfollows A. L. Jussieu closely. It is empirically true that it is possible to draw important characteristics from the flowers and the fruits, but in the background we still find the arbitrary assumption that generation is primary, an old scholastic concept. In animal systematics, however, the order of importance is: 1. nervous system, 2. respiratory system, 3. circulation, and the accompanying comment shows that choice is based only on the degree of generality. Lamarck assumes that if we consistently follow this weighting of characteristics we gradually approach the 'true order,' which is the order of coming into existence of the various taxa. Here, as many times elsewhere, Lamarck's conviction that his theory is correct is so strong that he fails to provide proof or further arguments. In all his theoretical writings there is this prophetic element of the assumed obviousness of his reasoning. In zoology, says Lamarck, it is mainly comparativeanatomy which will lead the way to the natural system. Briefly: the natural system is based upon the morphological organization of the living beings. Species (chapter III). Are species absolutely constant and as old as nature or have they changed, even though extremely slowly? The answer is not only importantfor zoology and botany, it is "essential for the history of our globe." Up to this point species were defined as each collection of similar individuals produced by other individuals like themselves. This is correct, because there is always this resemblance.Incorrect, however, remarksLamarck,is the secondary but almost universally accepted implication that the species are constant. This concept of constancy is pre-scientific. The first and best indication that species are not constant is their variability. The varieties and sorts so often described illustrate this variability; less informed biologists often assign to them erroneously the rank of species, thus confusing the issue. This variability is minimal under constant circumstances. By placing too much emphasis on such situations scientists have arrived at the dogma of special creation. The assumption that the living beings appeared not all at the same time but in a certain order does not necessarily detract from the religious dogma of an allpowerful Auteur de toutes choses. Lamarck was not an atheist. Guyenot (1941) pointed out correctly that in Lamarck'stime it was no longer dangerous to be a professed atheist. Lamarck's remarkson l'Auteur de toutes choses need therefore not be seen as a subterfuge; on the contrary, his religious position was much more that of a deist, and he even cites the famous dictum that one should not confuse the watch-makerwith the watch. Nature has autonomous laws but must have come into being through an Auteur supreme. Development of new forms and diversification in general by natural laws are characteristic of the world of living beings; without such TAXONVOLUME20 426 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions natural processes there would be a need for constant creation. From this it follows that Lamarck's cosmology presupposes an immanent finality. Development and diversification start afresh all the time: continuous spontaneous generation, ruled by laws of nature, is an essential feature of his theory of life. In his Discours of 1800 he stated: "In fact, the earth, especially near the surface in the water and even in the atmosphere at certain times and in certain climates, is populated in some way by animated molecules the organization of which, however simple, is sufficient for their existence." "These animalcules reproduce with a terrible fecundity..." The rate of reproduction of planktonic organisms has now been shown to be in general agreement with Lamarck's ideas; the idea of spontaneous generation, in this form, however, was effectively killed by Pasteur. With Lamarck it is a pure assumption; he never even tries to produce proof. Lamarck remarks that his views are not in conflict with the assumption of the existence of an 'infinite wisdom'; he, however, restricts himself to simple observation. Such observation shows that when our knowledge increases, our lines of separation between species (and genera) fade away. Nature is infinitely smoothly diversified. The existing animals themselves, however, and the same holds for plants, do not form a simple smooth series. They are arranged in a series with many branches, irregularly 'graded' but with no discontinuities, or at any rate with no such discontinuities in time. Present discontinuities may be due to the disapperance of species. Botany, which considers the other series of living objects, provides proof of this type of infinitely varied gradation and of the absences of really clearcut lines of separation. "How difficult is it to distinguish the species of the genera Lichen, Fucus, Carex, Poa, Piper, Euphorbia, Erica, Hieracium, Solanum, Geranium or Mimosa?" When these genera were set up, one knew only a small number of species which were easily distinguishable; but now that nearly all those gaps have been filled [sic!], our specific differences have become necessarily minute and insufficient." The examples show how well informed Lamarck was with respect to botany; present knowledge only reinforces his main argument. On the basis of this discussion of the nature of species Lamarck gradually develops his thesis of organismal development by active adaptation to changing environmental conditions. Even though it is debatable - to say the least whether this adaptation is 'active,' Lamarck's description of what happens is correct. "For example, if the seeds of a grass or any other plant at home in a humid meadow are transported in one way or another, first to the slope of a neighbouring hill, where the soil, though higher, is still humid enough to permit the plant to survive, and if later, after having lived there for many generations [sic!], the plant gradually reaches the dry and arid soil of a mountainous slope, and if the plant again survives through many generations, it will have changed ultimately to such an extent that the botanists will consider it a separate species." From this example it appears that Lamarck had a population concept in mind, and that his adaptations were of populations, not individuals. If species are really constant, they should not hybridize and produce fertile offspring. However, experience has shown that there are many hybrids between species and that some of them are fertile. This circumstance leads Lamarck to one of the most remarkable - and often overlooked - statements of the Philasophie zoologique: variation through hybridization can lead to speciation. "Truly, these singular pairings often produce no result, especially when they are very dissimilar; the offspring is then mostly sterile. When the disparity is less great, however, we know that these defaults will not occur. This procedure AUGUST 1971 427 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions alone is sufficient to create step by step varieties which become races and which, in time, constitute what we call species" (italics mine). This dynamic view of species and speciation has the following consequences: "1. All livingorganismsare productsof natureand comeinto being successivelyover a long period of time, 2. In its stridenaturehas begun, and beginsagaineach day, with the formationof the simplest organisms;those alone are formed directly and by means of spontaneousgeneration, 3. These elementarybeginningsof animaland vegetablelife went and go through a process of continuous differentiation, 4. The principleof growth,inherentin the simplestorganismshas led to processes of multiplicationand 5. With the help of generation... sufficient time, favorable circumstancesand changing environmentalconditionshave modifiedthe organsof the living bodies everywhereon earth ..., 6. Consequently... species have been formed gradually,imperceptiblyand successively;they can thereforenot be as old as natureitself." The circumstance that the plants and animals from ancient Egypt, observed and collected by the famous Napoleonic expedition, are identical with those of today does not impress Lamarck. He would have been surprised if it had been different! After all, the environments of ancient and modern Egypt are practically identical. The main point is: what are two or three thousand years in the immensity of geological time? This apparent stability will be mistaken as reality by the uninformed,because in general man judges everythingwith respect to himself. One has to look farther than a few generations. A striking example of change are our cultivated plants now taxonomically so far apart from their wild ancestors, an example in which Lamarck did not believe when he wrote his Encyclopedie. Lamarck'snew definition of species is: "each collection of similar individuals, which are perpetuated in that same state by generation as long as their environmental circumstancesdo not change sufficiently to make them change their habits, character and form." Fossil species. Many species known in fossil state are not known to be alive today. Lamarckadmits that he is not clear as to whether they are really extinct or whether they may still be found except for some large animals exterminated by man. What about those numerous fossil species of aquatic animals, not known to be alive now and for which annihilation by man seems improbable? They are probably the direct ancestors of related recent species and were thus not really exterminated,like the big Saurians. The main reason for Lamarck's uncertaintyis perhaps his geological uniformitarianism(actualism) which made him a strong opponent of the then still prevalent theory of one or more universal catastrophies, that strange offspring of the old story of the deluge. Such a catastrophe, says Lamarck, would provide a simple explanation for much that is still unclear, but there is ample evidence that this catastrophe existed only in the minds of men; there are no proofs in nature. Local catastrophies, such as volcanic eruptions, floods and earth quakes, are acceptable; but there is no proof of a universal catastrophe. On the contrary, in nature nothing is done abruptly, all changes are slow and gradual. Lamarck's belief in the continuity of the operation of the laws of nature evidently withholds him from recognizing clearly that climatic changes and other geological phenomena may also result in the destruction of life. He was sufficiently acquainted with the fossils in the Paris basin. His ideas might have been different TAXONVOLUME20 428 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions if he had had a better direct knowledge not only of tertiary remains but also of Silurian and Devonian palaeontology.The strong deductivetrend in Lamarck's thinking here shows one of its inherent weaknesses. Generalitieson animal, vegetable or mineral (chapterIV). It has been pointed out above that before the nineteenth century natural objects were classified as belonging to one of three kingdoms: animal, vegetable or mineral. Thus these objects were placed at a single level of comparison even though their origin was entirely different. Lamarck'sviews, as we have seen from his Encyclopedie and Hydrogdologie, were different. The primary division must be between organic, living objects, and inorganic, lifeless objects. The difference between animals and plants is also clear-cut. There are no 'zoophytes.' Animals can react to stimuli (can be 'irritated'), a property which is not at all the same as the 'sensitivity' of plants like Mimosa pudica, which is based on changes in turgor. Lamarck'sdefinitions of plants and animals are: Animals are living organized bodies whose parts always react to stimuli; nearly all digest food, and move either because of will-power (free or dependent) or because of outside stimuli. Plants are living organized bodies which are not truly irritable, which do not digest food and which do not move either voluntarily or because of true irritability. Tschulok(1937)has pointedout that in these definitions- which we find in several of Lamarck'sworks- plantsare actuallycharacterized as 'livingbeingswhich are not animals.'He tracesthis backto Lamarck's analyticaldisposition,so clearlydemonstrated by the Florefrangoise,in which the dichotomiesconsistmostlyof a positivestatement in contrastwith a negative.Tschuloksees this as comingvery close to the scalanaturae. The remarkaboutthe negativenatureof the definitionof plantsis well taken;Tschulok's furtherconclusion,however,that Lamarckhad not really evolved beyond scholastic reasoning,is too far-fetched.Lamarck'smain thesis is that plants and animalsare mutually independent lines of development of life. This is an original viewpoint, clearlygoing againstthe old idea of the chainof being. Lamarck, usually so refreshingly free from teleological speculations, hazards one in order to explain the fact that there are so many more species of animals than of plants. He states that there are, for instance, almost as many species of insects as there are species of plants. This statement is surprising at this early stage; it is essentially correct, although it now seems that the number of insects alone is probably twice as high as that of all plants combined. The enormous fecundity of animals might harm the general order, "if nature had not taken precautions [sic!] to keep this multiplication within certain bounds." In his Discours of 1804 Lamarck formulates it even more teleologically: "by means of these wise precautions of nature, order is maintained..." "Animals eat each other, except for those that feed on plants but which are themselves the prey of others." Essentially this is the modern concept of diversity through pest-pressure."Man alone ... seems able to multiply indefinitely ..." However, nature has endowed him with numerous passions which develop alas with his intelligence and which keep his numbers down. "Never will the earth be covered with the population that she could nourish. . . " After some doubts in the middle of the twentieth century, the Green Revolution has shown that this conclusion, if not the reasoning, may after all be true. The state of taxonomy in 1809 (chapter V). We have seen that Lamarck distinguishes between distribution, taxonomy, and classification, the simple 429 AUGUST1971 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions delimitation of taxa. These two methods have a different aim. Taxonomy (distribution generale) provides us with an order which must approach that of nature, that is the order in which the animals have made their appearance, and which can be deduced from their form-relationships. . . ? _- -. ?, ? .,?, I- .- . --.. ?" ,.."?-,*. *-,.. ., .- -. 1? * .-? 1s'-I*. ", ,- .II.*.'*1-. .:?p.I,-',. -,'".1 .-,,-. *1.-I? I I I .I I .I I- I I I- 1?..I I., .1I ..I .I? .I, I . . .? . .. -. .I: . ,- .I? -- ? . -I . , ? ., '*", -,I' ?- I . ?. - ,-.'-I1 ' ,. I .I " . . .I I," . ,I?, I1 .?"-1I . I?. I1 -. I .1I . ?. ? I I - , 1? -> I I ? I ? . ., I I . . ? I . 1. . I I. ?:i :?:?i;? :? *-".1'?'wl ,1. -*:i?i:??:i: I4 7 ?i:ii:??i:??:iii:i:??ij:?:: "::,:;:*::,?i:?::?:??:??i:::i:::i?i:.::::..# -?:iV ?:iiiiiii:, . ...i:.,. ...k ,.-? ,:?!::-,,, : ,".:::: , , .I?:: .,.:,:, ".,::::: ll. ..I 1. ..::! :?X ':. .-?... ?i?:??];:.:::::i:::i:::,?::::?::::::.:::::::::.:?::::::::?::?:::::?:?:?:::?:::i:::::?::??:::i: :I.:*,:! ",:;.,.., , , .I&,,jj::-, ..I?:.. ..X *,, ...::!,: I?. :?:,: :. I. ....... i *X :,::* :X:X ?*i*i:? i:?,?:?:i:?:i:i:?:i,??,;:;i:::?!: :;: *i*i*]*?:i:? j:.,F?.:j" X 15 N T 5 ...j:!: ,: "W :;:::::i:?:j:?,:;?:::::i:?:?:?;i;::i:i:??::::?:i:::::::?:j:?:::]:::?:::i.:::::::::::::::::.?.:]: i :,. : .? ?:: : : :;?: : ;: : i,: : : : : i: : :,. . : ?. . *;:.: ;:,?;: ?: ?: : : ?;:?:?:.?.?: : i ?i?i?]i :i i1i?:i :i:i:?i?: : i:?:?: ? H::i: :i,?:?:i:i: ,?: :i ,:,???:j:j:?,: i : : : ]?:???:????:?i??!:i ]????:?l: ?,:1 :? :.i : ]:i.? ?i I oiorli. e! :, ,.: , d ., :?-i:?:::i:i:?,i:i?i:::i,:,i:??.?:i?:::::.::::::i:?::::.:.*,::?.,.?::.,.?.I.?.?i*:?.?,??.,?..,. .? : kj: ?:] j: 7: j:?:j : j: ??::????!i??????ii?:??:?i .,i: ?:?:?: :?:i o -il vre, .i?i ? ?: :i?:i? ?* i:u 'i il?r .!$, -?'. :i :?i:?: : i ?. ?.;? ?:::i?::i??*i,::?:?:i:::??::::::i:i::;i:?:?:?:7?:j:?:?:?..:?.::::::::::,::i:::::::::::i,::i:;::;::::::i:?:::i:,P -. :? - .: -i;j: -"-: .:,.-: . ::::: ?: ] :.; : ,:? ' .:, .:- :. ,:. ., L I .: ?:; r. ?,: ; -:! ,:? .:.:. :.:.:. :.:.:?:.:,:.:,." -. . ,& w .,-j -: l : W -:!? I e -. I" : ": :;.! *? t l e ; I e -?* '." .", -l':?:.-*. -. : : ,:; : : .: .: .:i : .1-. * :, :?;, : - .:? ; * :; ,? # ;: :,. :.- ?,:;.,*: .:- .;: - :. ' ": . : l? 1, e .: . ., : :? :? -' ?: :?.' -:* i:* ?* i?: : i: ',...:::.;::::::.::::;:::::::?::::::?::::?;:::?::,;::::::?:::;:]:i::,i:i:i:i:i:?:i:i'!'?:?:', .' .: .* ..: -. .", "., . . * - , . -? . .- . - -, , ?: *- ., - .1 I . I I,,?- I ? -, .? .1I- .?- I -I1. - . -1 I ,1 . 1-? .1 .1 - ? * " 1 . I .1 I - ? I ? I 1 I.1 . .? I - :' 1.I I ? . , I. . . " ? I. "?,l.- . . . . - -. 1-., . ?",- . . I I - I II , ? I . I I I . ?l I I . ?, 1., - ?I ,? . . ? . ' - - . ? , . , - . . - . . .- "., 1. ? "I . -? II .- ? .- I I? . I - , 1 -. I 1.' I ?, - .l . I ? 1I. ? . I I - . 1: . .1I ".I I I ? I ..1" ? " " '.- -* ,* " , ., ? . . . . . " ' I ..- . . . - ,.I . , . ? . , ;' 1 , I . ?' 1?. -I I1 , P 1 ."., I.,-. ? I?. * I '. .? I ,1. . 1.I, I I -, I?. ? . ?II I. , I ?I, .I? ? . .- ? I,. . .? .I , ". I' ? 'I1. ? . .1:,?,- I :.?-' X?: :. :?!,:? -;?: - * 01 :-? : anirualix. :, ? : ? des ion at :x ;" : ? .; ?e" X : :?.: : :i: *.: : : : : :?: ?;:; I :i? . - .: .:? . 0 14 .4, " i zA. I d .,: "' ,;j:?:j:?:j:?:i:]:?:?:i:,?:?:i:?:?;:.?:i:i:i:?:?:j:?:?;j,?:?; :?:?:?:?:j:j:j:j:?:?:?:i: :i?j:?;i:; i:?:? : ., ?:., l :i?: i . :?;. ,. l t lu iv a n t ,:-s . .. I... II . I 11I I :::::: I i-,.,:i* ...:?:j:?:i:i:?:?ii?.??.????;::;:j??:]:?:?::??:j:?;::::i:', ...... :i::,?:i:?:::?:?:i:?:::?,?:::?+A ...',??:?:?:?:?:::]:j:j :?:::?;j:? I .1 :j:::?::::::? .:.: . . ? I I I I .1. ? 1, I ? I ' ,j:::?:j:j::;::??:?:j " ::?:?:?:?:?:?:j:?:j:j:::,:,; I"III?I',I I , II 1,. I.. . .*I ,?.. .. I: ,."I,*. a e oil, la fortnatl .ime des.,dn imaux '' w ."'j?;: j ? :?i?:i?i ?:? jiji? j?:i* i ?]??i?:i :?:???ii :?i]??:?:i?: : i: ?i?q 1 .J? .: ? * o AIR .,' :. A .:-* ? .,: :i ?i : i :i?i .,-:? :.- ile - I., i?fl I _. ,:?: .: : .: : ij: ,; i:i:?i?:?i??;?]: :??i??ii ??ii i??:i??:?i: i ??i?:?:,: :?:?:j:?:::i:?.,?j:j:?;??j:?:j??:::i:i:i:i:?:?.,?.:.:?::-,.i:?:j:j:j:?:::ii:i?:-i:;.:.i.::ii:i:::?,::i:i ,?:?:?:j:?:?:?.f. ??i?i?i???i???i?:?i???i.*'?i."i?i;??i;?;t ,$ j:j:j:j:?:?:?: : :i:i:?:?:?: :i:? :?:i ] ?i?i:?:?:? ?i? :? i:?i:i i:., 5.:* , .-" ? i? ? ? ?:? ? ? ? ? ?. I ? ? i? i:l iA k I . ::;::::::??:::::::::::??:::?ii?i:i:???ii::?:ii:?ii?:i.i:::?:?iiii:iii?i::??:i]:?'???*i]'?i?i?i??:?:?:::?:::?:?::;::?:::::::::::i:::::i:i::::?:::::::::::,?':.:: ,: : ?: 4* : *., :i?!i?j :?i :::::i:?:::::::!:::::::::::::;::::,:::::::::::::::::::::::,i:::i:?:?:j:?:?:j?i:?:i:::::??:::,?:.::::::::::::::::::i;i:j:?:i:::::i:::i:?:i:::i,?,?:]*:*?:?:j.:":?' ""i:i:i??:?:?:?:j:?:j:]:i:::i:i:i:::i:?:j:j:?:::i:i.*.,:::?:i:it i:,*::?:?:?:::::?:j:?:::]:::?:::?:::?:::i:?:i::?i'?,::?:::i?!:::::::.::;::::::::.,.::.:: : i*:, .: ;]!: ?: :j:?:j:?:?:?:?:j?: ?:j:j? `: : *: :, : :*:*:?. ,:?*ilil l:*?,?i,: : ?j:.,:?: ? ;,: : : : ::i::*?;:]:?: i::i,?li:??,i.-?"??i:iiI?,?il: ,?,? .1iI -:.X:- ? :? :? :? :?i:? :i?:? :? :? :?i'.? i? i? ? ?' ? ?:i? :i: ? :? ?i:?i? ?i ? ?i :; .: : :; : j: : j: . ': *?il :?.i: ?: : : ;:i .: : -.: -YX-.', :?;j:?:*:?;?:?:?:?:j:?:i::;i:i?::i:::::?::::;:?:::::::::: : .* "- -% ,: ! : serivraoo *-? - A r sGoP",arees : .:,*? subraincuses. I , ? ? :.*! :%:? '.- 2 &?, : : .? ?, : :? ? II I. ?-.?I4.a1, og I ?-... ,.,I?* ?i:?:1]:i?]?ii?i?]?ii??i??:?i?i?i:ii:ii:?ii::ii::;::?:::::i::::::;i;?:?:j:::i;i;::?;::::i::;::::i:??j :i?i. !??, ..I- -.?II :I?I.?I II .I WM ",*",:?:ii?i::??::;ii?il:?ii:??i?:??:??:i:::??::????iii??i??ii.?i::?j:j?::?::?:j:?'?,]:?: -I..???!?:?!????i??iii?iii?????ii?:?l: :. ?" .. 1. ... ...---x:j:j:j:j ......- i:?i:iii:..ft I., ??:',, I I II?,. :,.-.-11 ,:.: ..... :?:j?ji??i?:?i:i::??::j:?:,::;:::, ,?i;i?:?? :?::?:::::?:?:j:?:?:?:?:?;::i:?'7* j:j:j:?:j:j:j::?.j:j:j:i:`." ,."ii:::?::"??ft , -"'' ,:?' ..'-,,". *- Iklu, x -11 .....:: t 2 'I s ItM .,IE ID I.its A N I M A U . x .?.."-.?, ".1 ?? -??? ,Z.,.,.*' ::,??:?*?:?:?:::?:]:?:j:]:?:?:?:?:::?:?:i:::i:i:i:::!:?;??????? -?? -. f $ Z ---.??:i:? s J * v I ] : : :;: : : : : : ?,.: : .: : ,:?: ; ?: ;:i : ];?: :?:j:?:?:j? ' .: . . . . ..:,. :.. %,:. ". :., . ". : .: : : : : : : :!;: : : :?:. . .: ;: ".: : :???:?: :i :i:i: :i??:?j:j:?*?:?: ;!??ii:i . i??::??:i?ii:i??ii?i:?i????iiiii?:iiiiiiiiiii:iiiiiiiii:??i:iii?ii::ii'?", ::i:?:::i:::::i:i:i:?:?:::i:?:j?j:j:?:i:?:j:i;::?:j:?:?;?:j:j:j:j:j: :::::?:?:j:]:?:i:::i:::::i::::,:;?i:::i:i:;:??:?:i::;::???:?:j:?:?:?:?.:i :j:j:j:j: :j: :?:j:?;j:j:?: : ?:j:j? : ?j:?i: ? : ?:j:? '?: ?*?*?:?:::?:j:?:j:?*?'?.-. ::.?. :::?.-?.?:..":?:,-"*",, :j?j:j: :?:j:;: :j:?:j:? :?:? :j:?: :?:?: ,: :,. . . . . . ?-1-??.?.,?'- : : ?: : :?: : : :?: : : ?!? :",*-'?,?-.";"& -: .,- . . ,?. .,?:*:?.'*. ,.?.?. ?.".? -. . ,? ?:. ,. . '. . .:4 , ;i:::?:i:::i:?:::]:j?iii?i:::iii?i??i???:?i:iii:??i::?iii?,':?ii????iii???i? ji:? : :?:j:j:j:j: :?: :j:?:? .:, ,*:? . MI ". O F. '. . ? ? ;.: :.-" ".? ,i:.? - .: : b i! o . -o . l . :.u ; ? - '.?,'. -.',?_'. ? '. .:-, i ?:.6 .i:, :?, 6 :? i,. ,:? x x .l- 1 0 ?'o w - "',.-P.w . : : : ?: ?: ?: ?:: : : .: ? .:,i? " . ,-. .?. ;?..-,:,*,:::..*.:::::::::,. ::,?.,.::?l.-.:.,-.-,.-_?,:.".:.*.,::-,--::,..-??-??.,.?-. ,.,.,-. ?-.,.,. '?. . .,.,. . ::;:::::::::::::::i:?:i:::i:i;::]:???:::::?;::?lj:j:j:?:j;]:?:???:j?:::;::?:?:::?:?::??i?i?????:?:i:?:?i::?:?::::::t : : : : : : ,: : : : ? : :;: : : : ?:?:?:?:. ?:?: ? :j:?:?:j? : 1. AIMONTJ u ,.'.;0:owo,:'L "- '.? %X. '-.:; : :. ,? ::::;:::::;:::::::::,::..:::::?,?:?:?::,::i:i::.:::::::,.:::::::::.:,:.:.:::::::.:::.:.::: :?;j: j: j: ?: : :?:?:?:j:?:?:?;j:?:?:?:j:j:j: -'.I?t .'-".'"?-.T i? .-?- - - ?-., ?,?., ?. ?.-? .?.?t .......?.:,;.:?:::?i??i?i:i?::?:?:?i:??::???::i::;:,#,.''-.*,-.,?--.----,"-.,'""-?",*?',??'?-.-*. ,. % . :* :,*?: ., : ?: :.:.,: :. ;*?'-.?. :. :.:.:.:.?,:,:.?.: . !!??i:?:i?i:i??i:i.::?]?:,,.?ii:::il::?iii:?i:??:i???i?:i????????i:.?,:?:.?.:: ? i?i :?i :.i -?! :.,- ". :. :? -. -. -.,?",?."?', .?....,.?.-.?.:???.?.,::,i:?:j:?:?:::?:?:::i:i:?:::?'?:?:::?:?:?:::]:::]:?:],?::?:,?'....... :::::::?:::::::...::::i:liii:li:?i?:i?ii?i?:::?:?:??:?:?:?i??:?,???????????i?:? ;: -!: : :? : :, .1 4 -:.!:::::::::::::?:!::i::?i.,.,...::i:::::i:.."..:,ii:ii::::*??:i:::i:i:?:j:i::i:i:i,?:::i:i:?:?:1,?i:i:i:i:::i::.:i:::i:?:?:::i::,::i:i.::::::::?:::i:?:j:::::::i:i:?:?:::i:i:* ?i?i zi i; ? i ]? :!:i?:i :i ?:j:j;j:j:j:?;j:j:j:?:?:?;j;jI i*:?i :? ?:;: : : : : : : :?:?: :?. `?T . .:.:.:.:. *.'. ":.,.: :.,.: ,.,:,. .: : :. .?.: ., : 0 : la .", . . ??:: ?: :i:i:?: : i:?:?:.: : : :.:.: i:. : :,:?:i: : k:s4,'- ?, 'A ?.' li A?-., . ." ",':j:j:j:i:i: :i:i. .;. . . :?:?,?:?;i: ;: ?:j:?:?:l?j:?:j?j:j: : :?:?:i: :i:?:?'?,j:?:i:i: :?j:* ;:, :? %:* ,? . -?, l I'l 7 A . % T( It L S I . 1, :, :,; ,. :,? U IC -A - 1 .: : .1 . .. ,X * : - . .? V?: .15 l? I I ' l I - I .: 4 ? I . I .1 I ?,::` I:?-x-?..,. 111?;. .:;?::?:?;j::*, ::?:::::,: .?..' 1,??I ..." -?-.-?, .-.:.:::-:,??, .,.".,::: -'? ...-m ?.... ..: ::.,:!::: -:,::::?:7:1 I,1,,: ..... ..-..:.. .I:,? ...... ,.... 11 ... -.1I I .*/ I I..?,..:, ,;-:11 ':...'..;:??j:::;'...-%: ...:I I. . ., -.;:.. :. ?.1 ......"," ."? ?l,.-.-..-.-::..--,-. .:. .-::,:?:::;::? I., ':-l.I.... ..," i.--i?,i,? ?:?:,,.?', -.X,::?? ,.... ?'. "." '.*.?i,..'.:-:,;-'.',i .:"*,!X.?? .,.:X-:.-., ..?.,,.-.... .-.?.:.".--?--.,'?.. T# ,1. .1. ..-, -?... .-?V....II..., ::.:.*,.I .I II1.? " ,,:,::.: ... . ";;., ?-, I.l?'.".,,..-.',,'. '..*.'.'.* I .?I ? II, :,, l< I ..: .:.;,::::X,-?""'' -......... 1. -.1-11 -..??? ........ ...?---,`.-... .. **** ,. I....I' 'I',......,., . ?II :?:?:? ,IL " ' 'I :.. :-:,, ::::, -??-*.-I-:,:::? " .... ... ....???.?.??...:?:.:.,::.:::;:?:,:::?:,?::?::?":,,.....'.....".,.. ;.-, :" -.:?:*:j:j:?:j:j:j:j:j:j:j:j:?:?:i:i:i;:.:.:.:.?:,:.:.:.:. ........ ??... '", ?.?*.????'???????i;????????:??;:ii?::i:;::i::,:::.-:?:.:::: .:::;,i.:;::i?j:j:?,i:i::::,:,i::..":: ? ?::%::::'%? j:j:jxj$j:?;j?: , . " ? ,?.., . , ,*"i?ii::??i?iiii?:::??:?:??ii:?:???.?:i?::.,::,:.:::.?;,.,:. ?l ,.I., ,X-XI.,:? .:? .:,:,* -.?::::::::?:::?: :.:.:,.:-.,:, :, -]::?,j:::::::?:?:?:::j:?'.... ? ? ? :?:?:?:?:j:?:j:!:?:j , ? . I . ? 1. ..:.,.:.:.. ., " ,::..... .. ? ;.:!:.:?::::;::, ?-?'....... ., .:.:.:.:.: .-;.::;:.; 1? ,:., .7:* ? ?4 i*.i?ii:??i?:;i:?i?::;i:!ii?::i?::?:?:?: ?, 1:* "x ,.,:,:... -, . ? I .... ? ,.. .:iM. , ?_i; _ ......... ,"., ,,%,,,, ., -:,T:?::;:;:::i:iii;i?:: I I ?, :::3,:::::,:::;:::?:; -, -:p . ?... . ?... "I , : .. . -.?_:j:?:'* .., ........ I .A. -.??, ", ... ;,.?t ,,, ,.-.,,? . . I -,....,,-, .... ... ? ? ...... ::" ., , .... . . ? i:?:i?::?:?:i?: ,? ? ?X:: ,,, : -" . . ?i:;.".*,"?i.'. ...--::::;::-:-:,:::.i..;?i?.,i?:??????.,.. -:-:?:?,?.;: ,?:?]??:?i?:Ii??i?iii?? . .:?:,:,;:;::?.:,.,.,--.. I,. I . I? ? I . I-- I :'. .....::::, ...-1 ,-", ***-*,;:: :,.,", I. ,..,?-..,.... -....:?.'..?:: ". ::,:.x::;.,X.*..;? :'*-,:.: ....."'. -7:-..-.::!..*?.-:.::.,-,?,-*.?.!:--::?.::?.:?*?,,?::: -?, ....... " ?:?:j:?:???:j:?:?;?:j:j;?:?:j:?:?:?:?;j:?:?;?:, i?,.,,,,,"",:::::,..,-...,.,-."-,."..,,.,*."?.*.,*.,%,. ::--`-':,'5?,-l,...??':,,???*. -?-11. :,*" .:.,,.::::;::::;i:::::i:?:i,.!.: ..... ---....,....'..-.-.-.' -?,:",:,,::?.i.i??:??::?:j:??:,?:::i:?::i;?,?'?::::;,::: ....... ??,??,?-,.: ???:???::,:::?:?:::i:i:?:?:?:?:?:j:?;, :-.R. ?..... ..1" ;?:: ?I?... .,.-;::?$?:::j,?--.... -?. ,-'?' ...'-..'. -i??. :` ... ,-?C ...1 X.:::, ?-11 ?:]??Ii?.Iiil?]:i:? -".::,.-,.-.:;:,., -,"? .::.::-: -j?,* :" ::;:I',. .?. I-?I.,.:., ?? .,?:: ..-1-I.. -I:.?..I IIII.?.. ..:..I.11 .,::: .?:i:?;::::."?,?,?,?,?,?,?.,.,.'.'.'.'.'.......::::.,:. --:. ij.',]: *."??""... ?? .:.:.:.. ..-.;.? -.?-?:?'.-,.,--'.-..'-?.--.*."..I".-. ..."':;.?!:-:::::::::?? ??: -.;::,:::: ","? ,-.::; .I -.:.::::?::q.:::::?:,, .."F:,!,j:?:j::::::,??:j:? :::i:.:::::'' ,?? .... ?.:.:,:..7.:.:.:,:: :?:::?:::::::::::::???;:?.",:?::::,.,:.:*.,:: "I", :: ?::.,;,:: ." ,W. ...X ... `:,:,;j:?:?:::j:::?:?:j:j:?: ?;::.', :j:j ?:::?;?.;;?::i::::::?;???:;;??x*:.'?? --.**.,'.?'...-:?? ..':]?j::!::?::?" I,,?.,?? .... :: X .:!.: ... :::? :!::;:;::::::?:!:??:?;: ,,.:., X :::;:::..., ..::::::::::::,., ..,. ',...:.:::: .i??'*?i?:.iii?:.?; :. . :.,::.,..,...:.:::. I11 ,;.".".".::?:i:?",.,i:i: ?, ?::: .................... ..-".-'.'..... '!: ..,.1... ?,.,?i?;i:?:?:?:i::i:i:i::::*.,.,?:j:?::.:.,.,.:..,.:::.:::: *::-: .:j:j:?$j: "I ..?11 .I., .11-?:,?:::,:-::... X::::::t;:: ...: :: ........... :;::::;:::::::??.:: : ??]:?.'.-.'*.'?i:;.. ..:::... -.II.:,??I?-I?:. ,.:,:!?:,::::??:::::?:.:*;.,:::::.*::::..:::: I: !::::.., .. ...I11?1. ..-,:,. 1, ... ... -:;.? ?III. I",I;. II,I,... ': :::!::!, ...... ..*...... :::: ?:?:?:.:?:::i::::::.:::::,?:::!::?::: ,, :.:.:.:l,:,X ....:-.:::: ':i::?:::::i::::::::j::::::::i::::i:::?:;i: . ..:.:.:.:.,.. ...,:.?,:?::?:?:.:?:.:?:!?::,::.::.;:.:.,:.:.::.:.*:.: .. I 1 ??. Z ::X :::". : -::X I... .. : ':? . . .. . ? I :?????:?:?:?ii,?:i:?:i? ... 11 ... I .... .... . . ....... ?... ? :::::??:?;j:?:?:jl I .:::-;-:I ? ? * ... **" * ::.:.: I ...?,;:;:,:-:;,.-:-:-:,?.:.:.::?:,:.::.:: I .. I.:: :;:? ::, ::::::::: ?.,- I.I.. . , 11 ?.,::: -:1. - I III I ??:` ,i?:ii:iii?iii:i:i??i,..?:i:::::?:i: ..:!:*,:??,.,,? ::::: ." .?,, I.1I I11III I, -. ?II. ?l1-1 ,.:: .-...." *,..:" ...":: .;.,.......,..:.,:.:.,.:.:::::::,:.,::::::: :::::::?:::?:,::::,;,:...,..,,.,........,?.:?*:*::,:,:: ::.!::??:?:?:? ?.1 :i??i?? ?i:? I., ::;,:::::::: ::.:.:.:,:: I ;:.?i?",' .."...*i? i* ::.? ,..:ii ,?I..:.-:-X :: . .:.. ........ " -:,:.?-!?;? ::7,.-.::r::::.-.:.:.' :,;;:. :i-?i:::i?i?':]?::ii:i:ii?iii?:: . :?:?:::?:?::?i::;::?:j:?:::?:i:::::]:j:?:i:i::;::::i:?:j:?:::??:?'ii..:....:??::i??:?? ?? :: :::I:...:.;.:,:. . ... I ..:. . X ??; X ?-: ? . I '.::?;,. I ? : :. I I I . : 1 1 .,..,..:::::::::::::::..:.:.:.:.:. ..: ....... :.:.::::.:::::::::, ? I ....:. I I I . I.. :*.,*:::? I .1 . 1. .:, , ::?::?::i:?;j:?ii?:?i????]?i??ii??]???i?r . . ?*..,?:? ;?:: ,?i?:?:?:i:i:?:i:i:?:i:?:i:?::i::,i,::::::?;ii:i,:,::? : ...... ?, ,.."i?:N I 1. I I . I'l l I :.;?:??j:?:j?::?:j???:.i:? -:,I-II?:.,-... -, ... ...'' "x ''.. . I ?, -:::: - ,, :: ,.% ?": ..., .?? I-??.. ?I,II,? .: -;.:, x?:?: :.:......... .?:. , ::;:?:::::j:j i?:?:j:???:]:j:?:i:i:ii:::?:?,?,ii:ii*i:i:::::?,.i:?:j:j'j:?:i::i:ii??!?:?:i:i??????i :,:.::-X?:.:,.::i:?:!??:??i:i:i:'?:?:ii:i:i:i?:;?.:i:::::i:::: ...... .... .... .....7 + :.11:,?? ::! .:. :.?II:; ,,?:.",:-:: .:,.:., ..',?:::::::::;*::::;::..::::i:i:s?:::::?:::::::::::?::?::?::.::::::::;::!.: I-I-I ?. -,X .::,.:;.: Ib-:I1. I?.I-.1?, ..; ' .jjjjjj?? :::::::?:::::,::::::??:::::::::::::::::::::::::;; .1 .-I-......... ,.,-..::::?, ,.... -......" :i:??,::*i:i:??:?:i::??:::::i:i:i:i:i:i:?:?:::i:j ... -?, ..,', - .i?ii,,-?:: :?:?:.:.?.:.::,::::::.::,.::::::: .,. -,7.:7:::::!.*.:,:*:.X ;-?....,.,::::;::::::?;::::: I III? ,.I..I .,Z,I .... ,., I,::1? ':;,.:-:-:::*,,,?,',.:;: .,:? .,.,:::v:::,..:.:l "........ .... ,??,?, ....-.,-,.,*:j:j,::?:j:j:::: I . . -I1 I?l .".11 .,.. -.. .,. I .11 -.,:,'.,..:.,. ?,:,.."?'-.' i*:*i-i :`:' .,...,.,.,, :::::::,:i:: ..I,,- ;;?-,-.,'?-.'-'..-'-,:."..,.,i:i:i:-,i; -.:.:.:.:.:?:.:.:.:.:,:.:.:.::::::::::::::::::;:::j:j:j:?:::j'':j:?:::?:j:".,-* .--'.--... :::::i?j:j:i?:::::ii::ii:iii]?i:??i:]:i;iii:??i????iii??ii?i????iii???ii: : : :i ?: :?:. :., .*. :;., ., .: , i :i : :i i :?: :?:j:?:j:j:?:j :-.'.:". :-?x. " :'- .- ;. .: .,: ;.-:;. ,-. :- :,-. x : :: 1 : :- .- :, :. .<: :-,.*-l: ,. -: . :?:.:. :.;.i ?i?: ]?: i:i ?;i:?i ??:i: :?:?: .: ." : : : ,-: ?: : . , :.; :. ;: : ?; ,: :X ,: "?X: ?'.- ,-?: : ? j-: :j : .: ,:. " .? :i -*M ' ? ., -X ,:? : ?: : ? :! ?! :" ?' : ?:; : ,.: ;: -:,. :? : : : : ;:? ', :! . . I. :. : ?. .: !?.-:, I?: ,: : I , .:?*: :- ,.: ,:* : I. ? ": ! ,: , ,: o , , ? '. 1. 1" I 1 ? :jX , 1- ,.?: : I .? I., . : . ? I : .1 X :. . `:" .:?:-:..-,.," 1^,.? ?' ? ?.,:,i,?:i:?:i:i:i:ii::;i:?:?::,?i:'i?:'?:??:ii:"iii:??? -.1.-.?. ...... - ?.? -17!10" ";;:::::::::::: ;,..::".,,:?*,.:??:::: FIc. 5 TAXONVOLUME20 430 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions Classification, however, establishes dividing lines in the general series of beings, rest areas for our imagination, enabling us to identify the animal or plant. The criteria for the delimitation of taxa are conventions such as, for instance: "each class must comprise animals with a particularsystem of organization." Another important question facing Lamarck was that of a taxonomy with reticulate diversification (the idea of taxa as countries on a map of the earth) or one with an essentially linear arrangement.In his discussion Lamarck does not really give any argumentsagainst the 'reticulatetaxonomy' such as we have found for instance with Linnaeus. In his Systame of 1801 he says "This idea, which has seemed sublime to some modern authors who have badly studied nature, is an error which, undoubtedly, will disappear as soon as we have a more profound and more general knowledge of the organization of living bodies." In the Philosophie zoologique he simply states that the development of life is one of increasingly complex organization, and that because of this the main higher taxa (les masses principales) are to be arranged in a single series. We have seen that his ideas on speciation allowed for 'side branches'; the series is not one of species, but, for instance, of classes. In later years Lamarck abandoned this simplistic taxonomy. In the supplement to his introductory volume of the Histoire naturelle des animaux sans vertabres(1815) he presents an "ordre presume de la formation des animaux offrant deux series separees, subrameuses"which really amounts to a complete breakdown of the single series concept (see fig. 5). He adds in a note that the typographical devices were too primitive to express his ideas of divergent lateral branching. In the Philosophie zoologique, however, Lamarck still clings to his idea of a ladder of main taxa, a left-over from the past. His opposition to a 'reticulate taxonomy' stemmed from his conviction that the actual evolution had taken a main single course. On this basis of organic evolution it is obviously also possible to come to a system of branches of the 'tree-type'; a reticulate taxonomy, however, could never reflect evolutionary development and divergence. Lamarck's rejection of the latter was therefore a matter of principle: taxonomy had to reflect evolution. In his discussion of the process of organic evolution Lamarck touches on the problem of 'orth genesis' of organs (of which he denies the existence), on the phenomenon that can be called 'Spezialisationskreuzung'and on similar extremely interesting evolutionary processes which he discusses on the basis of pure deduction, and all avant-la-lettre.They all stem from his concept of one mainstream of increasing complexity. Each class is by itself a series, a part of that mainstream. For the animal kingdom Lamarck recognizes fourteen classes in a single series. He admits that in botany the situation is more difficult but hle thinks that with the application of characteristics derived from the seed and the fructification the final order is now in sight. One main question remains for both taxonomies: do we go from complex to simplex, or vice-versa? The mechanism of evolution (chapter VII). Before elaborating his own taxonomy, Lamarckdiscusses what is now usually known as 'Lamarckism':evolution through active adaptation to changed or changing environmental conditions, that is, inheritance of acquired characteristics. Although Lamarck as a thinker was highly original and independent, he had nevertheless predecessors who evidently influenced his thinking. The general idea of transformism, derived from general systematics, had first been advanced AUGUST 1971 431 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions by Buffon. Lamarck knew Buffon quite well and admired his career as well as his writings. The influence of Buffon's general ideas on organic descent on Lamarck must be admitted even though he refers only sparingly to his great predecessor.With respect to the mechanismof evolution it is likely that Lamarck was influenced by a now little known Paris physician, Cabanis*, who published twelve papers on Rapports du physique et du moral between 1790 and 1796 (published in book form in 1802). Herv6 (1905) has drawn attention to the Cabanis writings: in the midst of a mass of irrelevant medical detail one finds some striking ideas. Cabanis argues that life originates by continuous spontaneous creation (not new), but that organisms evolve further "under the influence of the climate and all other physical circumstances. . . " Domestication results in the acquisition of new characteristics and new habits. These dispositions acquises are transmitted from mother to children and are later maintained even in the absence of the original causes. Cabanis furthermore maintains that an organ can be strengthenedor weakened by the environment, a change which can also become hereditary; this can be seen as the first formulation of the law of use and non-use. Cabanis is therefore a convinced evolutionist. Fossil remains were his evidence that species had not remained constant. "It is not at all proven that our present species are the same as when originally formed"; ... they "carry the imprint of the climate in which they live, of the food which they use, and of the habits... " which they have acquired. According to Roule (1927), these publications, coming from a confirmed atheist, stimulated much discussion in the years before 1800, and it is clear that Lamarck must have known of this. Many of his ideas were therefore not really new: the general concept of evolution can be found with Buffon; the idea of the gradation of organization, the ladder of nature, was obviously very old and, in the form proposed by Lamarck, previously expressed by authors like Adanson and Bonnet; his ideas on the mechanism of evolution are akin to those of Cabanis. These ideas were in the air, but this does not detract from Lamarck's importance; by systematically bringing them together and by working them out in great detail he became the real founder of transformism. It is here not the place to analyze Lamarck'sideas on the actual mechanism of evolution (see e.g. Guenot 1941). A brief survey and some general remarks may suffice, because this aspect of Lamarck'swork had had a less direct impact on systematics. The main points have been touched upon above: changing environmentalconditions cause hereditary adaptations, creating new needs and new possibilities which are followed up by nature; it is not the organization of the living body which permits it to live in a certain habitat, it is the habitat which influences organisms to adapt themselves. The question whether this applies to populations or to individuals is usually not discussed. Lamarck maintains that this is not a theory, but the statement of a positive fact. There is no doubt that adaptationexists and that each species has its niche; however, he does not really show that the niche makes the species. Many of Lamarck's remarks on the actual mechanism are understandable only in the light of the primitive knowledge of zoological and botanical physiology of his time. They have, however, only an incidental value. Animals acquire new habits under new circumstances.Plants, however, have no habits; they change under the influence of nutrition. A gap appears gradually between individuals of a plant species which are continually well nourished * Georges Cabanis, 1757-1808, French physician and philosopher, forceful promoter of empiricism and positivism in philosophy. TAXONVOLUME20 432 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions and others which are not. "How many examples could I cite with respect to animals and plants which would prove the correctness of this thesis," says Lamarck, but as usual the examples do not follow. He then gives the abovementioned example of the grass species coming from a wet meadow and adapting itself to a dry mountainside, significantly evoking many generations and leaving out of account the question of 'active' or 'selective'. He also states now (contraryto what he said in the Encyclopedie) that our present wheat must have been brought to its present state by man because it does not occur in nature. "Where do we find in nature our cabbage, lettuce, etc., in the form in which we grow them in our gardens?" Domestication of animals has also resulted in considerable changes. The example of Ranunculus aquatilis is also interesting; the submerged leaves are different from those floating on the surface of the water. However, no floating leaves appear if a plant happens to grow in a humid environment which is not under water. The plant then becomes "Ranunculushederaceus, which the botanists regard as a species . . . " The latter is certainly true, but we know that the land form of Ranunculus aquatilis definitely belongs to another species. The two Lamarckian laws are (translation Packard 1901): "First law. In every animal which has not exceeded the term of its development the more frequent and sustained use of any organ gradually strengthens this organ, develops and enlarges it, and gives it a strength proportioned to the length of time of such use; while the constant lack of use of such an organ imperceptibly weakens it, causes it to become reduced, progressively diminishes its faculties and ends in its disappearance." "Second law. Everything which nature caused individuals to acquire or to lose by the influence of the circumstances to which their race may be for a long time exposed, and consequently by the influence of the predominant use of such an organ, or by that of the constant lack of use of such part, it preserves by heredity [generation] and passes on to the new individuals which descend from it, provided that the changes thus acquired are common to both sexes, or to those that have given origin to those new individuals." Lamarck's prophetic inclination, making him look beyond direct proof, is again apparent in his first commentary on these laws: "These are two constant truths which can be ignored only by those who have never observednor followed nature in its processes . . . " And again: "It is not that the organs ... have made possible the habits and particular functions, but rather that the habits, the way of life and the circumstanceshave ... in time determined the form of the animal body, the number and character of its organs, in short, its characteristics." Lamarck illustrates his theory with many now well known examples, such as that of the long neck of the giraffe and the features of the Kangaroo. He sums up: "Conclusion so far admitted: nature (or its Author), when creating the animals, has foreseen all possible circumstances in which they have to live, and has given each species a constant organization as well as a definite and invariable form of its parts which force each species to live in the places and climates where we find them and to preserve their habits." "My own conclusion: nature, producing successively all species of animals, beginning with the simplest and finishing its work with the most perfect, has gradually complicated their organization; these animals distribute themselves over all habitable regions of the world; each species has been submitted to the influence of the circumstances which it has encountered and, from this it 433 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions has acquired the habits which we now know and the modifications in their parts which we now observe." Lamarck's taxonomy (chapter VIII). After stating that animals and plants can be arranged in a series of decreasing or increasing complexity of organization, and then expressinghis theory on the actual mechanismby which increasing complexity takes place, Lamarck rounds off the first part of the Philosophie zoologique, dealing with general systematics,with a survey of his own taxonomy reflecting 'the natural order.' The importance of taxonomy does not lie simply in the grouping of living beings under 'species,' 'genera' and 'classes.' The taxonomy (the actual system) must "at the same time offer the most favorable means to study nature and to discover its progress, processes and laws." This view of systematics as a general framework for biological science is another very modern feature in Lamarck'swork. The actual elaboration of the taxonomy need not be given here. Last years: an aftermath too long (1809-1829) The main works of Lamarck'slast years are his Histoire naturelledes animaux sans vertebres(1815-1822, 7 volumes) and the Systeme analytique des connaissances positives de l'homme (1820). The introductionto the first work was for a long time the most widely distributed theoretical work on zoological taxonomy mainly because it was soon published in a second edition which, unlike the Philosophie zoologique, quickly found its way to many libraries. Little new is added to what had been said five years earlier in the first part of the Philosophie. The book is well organized and in excellent style; it lacks the many repetitions of the Philosophie and thus becomes Lamarck'sbest theoretical biological treatise, reflecting his whole spiritual development.Foremost perhaps is the insistence on positive facts, facts that can be observed by any unbiased witness and which are described in 'physical' terms. In another excellently written essay of this period, the article Espacein the Nouveau dictionnaire d'histoire naturelle (1817) Lamarck states: "I have said it often enough in my works, and I have no fear of being mistaken: we acquire positive knowledge only by observation, and we can observe only nature, its objects, their qualities, their characteristics and consequently only the products of the acts of nature." In this article, as well as in the Histoire naturelle and the Systeme analytique, we find again many a priori statements, based on deductive reasoning, which occasionally sound scholastic. The imaginative generalisations which proved so fruitful are stated more succinctly than before. The untenable extrapolations from generally recognized phenomena, such as the deduction of inheritance of acquired characteristicsfrom the phenomenonof adaptation, are still present. Throughout these works of uneven genius runs the red line of evolutionary thinking, of systematic reform and of a positivist approach to nature, admitting no metaphysics in the description of its laws and structure. The "fundamental principles" formulated in the introduction (pp. 11-12) are (freely translated): First principle: Each fact or phenomenon observed is essentially physical . . ., Second principle: Each movement or change, each acting force ... observed in physical bodies is necessarily determined by mechanical causes, Third principle: Each fact or phenomenonobserved in a living organism is at the same time a physicalphenomenonand a productof organization, Fourth principle: There is no such thing as general living matter; each living body has a specific organisation..." From such principles, but also from the Systhme analytique des connaissances 434 TAXON VOLUME 20 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions positives [sic] de l'homme (1820), it is clear that Lamarck is essentially a scientific positivist. He could, of course, not yet share (and it remains doubtful that he would have done so) Auguste Comte's religious positivism. Lamarck's more elementary scientific positivism is the direct outcome of a movement begun by the Encyclopedists to free science from metaphysical bonds; he has almost achieved this full freedom. It is possible to indicate scholastic and teleological elements in his many writings, but the main tenor of his work is empirical as far as the descriptive part is concerned, and positivist as far as his theoretical deductions go. Observed facts are the only basis for science. Scientific concepts should be free from metaphysics. In this way he could ultimately break the bonds of the past as embodied in, for instance, the static chain of being, the fixity of species, and essentialist classification. In the introductionto the Philosophie Lamarckhad already stated (p. XXII) "Only the facts that man can observe can be positive truths on which he can solidly count, not the conclusion which he draws from them..." In the Systeme (1820) he states "observation being the basis of my work, it seems difficult to me to find a better one..." The only yardstick is nature, not man. In this way Lamarck could achieve insight into the immensity of geological time and into the necessity of acknowledging organic evolution. In the Systeme he says also, "it is an indisputable fact that there is no absolute rest in the physical world, no absence of movement, no immutable matter with infinite rather than relative stability. . . " This sense of relativity, which he extends to human knowledge as well, is a typical modern romantic sentiment. Lamarck's later works are no longer footnotes to Plato; they are a breakthrough towards independent nineteenth century science. This feeling of relativity also leads to a sense of glory of the imperfect. The Platonic tradition, the essentialist and static view of nature, tended towards the perfect; the principle of plenitude itself was an expression of the perfection of this world. As long as species were static, definable by their essence, as long as they had all been created at the beginning, there was no room for imperfection. Organic evolution presupposes imperfection and increasing diversification: two typical characteristics of the romantic movement. Thus seen, Lamarck is one of the most remarkable men who helped bring about that great transformation of ideas and values which characterized the early nineteenth century. Static diversity disappeared in favour of dynamic diversification, be it in biology or in the minds of men themselves. Lamarck's later days must have been lonely. His far-reaching ideas did not endear him to the establishment of the day. The incident of 1809, when Napoleon snubbed him when he tried to present him with a copy of the Philosophie zoologique, was probably the beginning of a social isolation which in later years became heightened by the failure of his eyesight. Landrieu (1909) draws our attention to a contemporarytestimony by SainteBeuve in his novel Volupte of 1834. His hero follows Lamarck's course at the Museum and states that he fully realized the hypothetical paradoxes and the contradictions within more positive [sic] and advanced systems. However, he was touched by the "ton passionne et presque douloureux qui s'y melait a la science." Sainte-Beuve proves to have very well understood the essence of Lamarck's teaching, because he goes on: "His concept of things had much simplicity and sadness. He constructed the world with the smallest number of elements, the smallest number of crises and the longest possible duration. A long blind patience, such was his genius of the Universe. Once that mysterious power of life was admitted, as small and as elementary as possible, he saw it 435 AUGUST 1971 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions develop little by little in time, the mute urge, the sole habit in diverse environments gave birth at long last to the organs in opposition to the constant power of nature to destroy them..." The conspiracy of silence "More than half a century had to elapse before the philosophical concepts of Lamarckcould emerge from obscurity; even during the lifetime of their author, there was around them the conspiracy of silence" (Landrieu 1909). Why this conspiracy? The most simple answer is that many of Lamarck'sideas which we now retracewith respect were far ahead of their time. The scientific world of the conservative Empire and the reactionary Restoration was simply not ready for the general idea of organic evolution or the concept of geological time as advanced by Lamarck.However, such an answer is far too simplistic; Lamarck, with all his merits and demerits, was his own worst enemy. The idea of organic evolution was really advanced as a theoretical deduction. Lamarck had no empirical proof available. The existence of now extinct fossil species could also be explained differently, the more so since Lamarckhimself had no clear ideas about this aspect of evolution. After living in an anthropocentric world with a two-thousand-year-oldtradition of timeless essentials and man as the measure of all things, it was difficult to switch over to the concept of an immensity of time, millions and millions of years, in which man had played no r8le. Even with a great permissiveness towards atheism, religious dogmas still beclouded the minds of many scientists. Besides, Lamarck's theoretical weaknesses were very apparent to his contemporaries; at a time when the 'pneumatic' theory of chemistry was sweeping away with ample experimental proof the last vestiges of the phlogiston theory, Lamarck maintained through all his writings his theory of the central function of fire, which was still akin to phlogiston. At a time when scientific meteorology, and especially metereological prediction, was still in principle impossible because of the absence of a system of instantaneous communication over large areas, Lamarck tried to predict the weather, and published long range forecasts which simply had to result in discrediting him in the eyes of his contemporaries, and not only of Napoleon. In a period of increasing empiricism which cast more and more doubt on the existence of spontaneous generation, Lamarck maintained such a thesis as the starting point of his theory of evolution. With ample evidence of the direct volcanic origin of many minerals and rocks, as presented for instance by Hutton, Lamarckmaintained that all minerals were of organic origin. When typological thinking and advancing comparative anatomy clearly showed that species and genera had multiple 'reticulate' relationships and could not always be conveniently placed between a simple and a more complex neighbour, Lamarckinsisted, at any rate until 1809, upon a linear gradation of all organic beings, admitting only that the animal and plant series were independent.When drawing attention to the indisputable phenomenon of adaptation, Lamarck chose as an explanation a mechanism (genetic changes induced by environmental conditions) which too easily lent itself to caricature. Pointing out the phenomenon of variation, Lamarck explained it by invoking a directing force for which he had no proof at all. In view of all this, it is not so hard to understandwhy Lamarck'scontemporaries found it difficult or impossible to separate chaff from grain. This was the more difficult because that process would have resulted in upsetting traditional belief in creation, as well as many lovingly treasured theories. Apart from Lamarck himself, however, there was certainly another major 436 TAXON VOLUME This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions 20 set of circumstances contributing to the conspiracy of silence. To summarize that set of circumstances by citing one name is admittedly an oversimplification, but it spotlights the attitude of the Establishmentwith respect to Lamarck in his later years. The name is Cuvier. Georges Cuvier (1769-1832) was one of the great promoters of the natural system in zoology. His extensive comparative morphological studies, duly extended to fossil species, led him to a typological approach in his classification of the animal kingdom (1812). This system is based on four essentially different plans of anatomical structure ("quatre plans generaux... d'apres lesquels tous les animaux semblent avoir ete modeles... ") molluscs, radiates, articulates and vertebrates. Cuvier was a convinced believer in divine intervention in the world of living beings and a steadfast defender of neoplatonic idealism in biology. His idea of geological time was primitive, his view of geological history was heavily tainted with the ancient belief in catastrophes. Geological unitarianism, explaining geological events by means of the same causes in the past as we see in operation today, was as foreign to his mind as the idea of a gradual evolution of life. His view of the history of life on earth was, "if anything, 'extinctive,' eliminating by catastrophe,and not 'progressive,'creating (through God) new and higher creatures as an aftermath of catastrophe." (Coleman 1964). Cuvier rejected the idea of an animal series, whether in time or as an abstract notion, in favour of multivariate relationships between taxa. "Cuvier... seems to have been constitutionally unable to support or to appreciate the basic idea of change" (Coleman 1964), a state of mind which is characteristic of any Establishment. His comparative anatomical research contributed greatly to the perfection of the natural classification in zoology; his theoretical background, however, was thorougly eighteenth century and still imbued with many a priori assumptions. He described his own r81e in zoology in the preface to his Regne animal as follows (translationby Lurie in Agassiz, ed. 1962): I had then, and this took me considerable time, I had to make anatomy and zoology, dissections and classifications move abreast of each other; to search in my first observations on organization for better arrangements through which to arrive at new observations, and to use again these observations to perfect the arrangements; finally to take from this mutual fertilization between the two sciences, the one by the other a zoological system appropriate as an introduction and guide in the field of anatomy, and a body of anatomical doctrine appropriate for the development and explication of the zoological system." Cuvier was, therefore, in almost all respects the antipode of Lamarck. It is not the place here to discuss the ideas of this great zoologist, because they had relatively little influence on the stream of ideas in botany. Theoretically Cuvier was very much a man of the past although his descriptive work greatly advanced the progress of zoological systematics. Interesting and accurate analyses of Cuvier's biological thought will be found in Daudin (1926) and Coleman (1964). An extremely astute comparison between the characters of Haller and Cuvier was made by A. P. de Candolle (1862). Cuvier's position in the natural sciences during the Empire and the Restoration in France was extremely powerful through a variety of circumstances which cannot be analysed here. He was as well seen by Napoleon as by Louis XVIII, he held most of the key positions in biology of the time and published successfully; he thus embodied the established order. His authority was almost generally acknowledged. Scientifically he succeeded in demonstrating by means of his comparative anatomy the existence of a succession of different animals AUGUST1971 437 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions during the history of the earth, but he failed to appreciate its significance with respect to the views forwarded by Buffon, Erasmus Darwin and Lamarck on transformism. Cuvier became the prototype of the antitransformist even when he had at hand, in greater quantity than any biologist before him, the material which still most strongly supports the theory of organic evolution. His neoplatonic and actually essentialist theoretical explanation of this evidence was expressed in his type-concept - essentially teleological - and in his firm stand with respect to the fixity and reality of species. No wonder that Lamarck found in Cuvier his most formidable opponent. During Lamarck's lifetime Cuvier attacked his transformist theory only obliquely, in 1812, in a general introductionto his Recherchessur les ossemens fossiles de quadrupedes. He attacks the idea that fossil species might be the ancestors of different recent species by a reference to the animals known from ancient Egypt which differ in no way from those of today. Apart from this Cuvier acknowledgedthe value of much of Lamarck's critical taxonomic work in the invertebratesby adopting part of his classification. The real blow came, however, in the Ploge de M. de Lamarckwritten by Cuvier but read only after his own death at the Academie des Sciences on 26 November 1832. Cuvier's Ploges are curious documents. His Ploge of Adanson, for instance, (see Nicolas 1963) drew a caricature of the latter's contributions to science; it helped to strengthen another conspiracy of silence: that around Adanson. The Ploge de M. de Lamarck had exactly this effect with respect to Lamarck. It inaugurated (Martins in Landrieu 1909) "that genre of eloquence which would later be known under the non-academicname of 'reintements' [cuttingup]." There are undoubtedly several positive elements of appraisal especially with respect to Lamarck's descriptive work on evertebrates, but on the whole the dloge is unduly and unusually critical and often sarcastic. Statements like "moins de severite dans le discernement de l'&vidence,""conceptions fantastiques," "... *vastes edifices sur des bases imaginaires semblables a ces palais enchantes de nos vieux romans..." are abundant. With respect to Lamarck's - after all so admirably correct - estimate of geological time Cuvier remarks: ... it is always time which is one of the necessary factors of all things; that limitless time, which plays such a great r81e in the religion of the Magi, plays a no less important part in all of Lamarck's physics and it was on that time that he rested to quiet his own doubts and to reply to all objections by his readers." Cuvier's attitude and that of his 'disciples' is certainly in part responsible for the conspiracy of silence. However, there was some contemporaryappreciation. We have seen above the poetical appraisal by Sainte-Beuve's alter ego, d'Amaury, in Volupte'(1834). Etienne Geoffroy Saint-Hilaire, another and later adversary of Cuvier, stood up for some of Lamarck's ideas, in particular on variability, the non-constancy of species and transformism in general, thus showing that it was possible for a contemporary scientist to recognize the valid theses in Lamarck'swritings. Another great classical French author was deeply influenced by Lamarck., although by other aspects of his work; Balzac's Comedie humaine is based on an analogy between society and nature. If circumstances mould animals, society may also mould man genetically. Balzac's thesis is strangely reminiscent of the reasoning underlying a more recent revival of such ideas on the possibility of changing organisms - and man - through society, a revival which will remain connected with the names Lysenko and Stalin. The conspiracy of silence around Lamarck was not absolute. True, his main 438 TAXON This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions VOLUME 20 thesis that classification should reflect descent was not immediately accepted. On the contrary, the nineteenth century first witnessed a great outburst of more refined typological work for which the question of evolution remained of secondary importance. The controversy on the stability of species remained alive, although under the influence of Cuvier and his disciples the parti pris against mutability became stronger rather than weaker. During the last years of his life Lamarck found little open support either in France or abroad. The direct influence of his ideas on the systematics of his own time was perhaps small but these ideas have ultimately proved to represent the birth of biology. Appendix 1. Translation of Lamarck's letter to Pancoucke (publisher of the Encyclopddie methodique) of 30 December 1790. "Sir. I send you herewith the plan of the work which I propose to write with respect to the botanical part - which you have kindly entrusted to me - of the new systematically arranged encyclopedia. This plan has been drawn up with two objects in mind: [1] to bring about the most essential changes necessary in the articles of the old encyclopedia so as to make them truly useful, and [2] to write them 'in accordance with the spirit and the ideas which should be the rule in all special dictionaries of the Encyclopddie m6thodique. I must admit, Sir, that when you charged me with the compilation of the botanical dictionary, I had hoped to find in the old encyclopedia, which I did not know at that time [sic!], a considerable amount of good material which would have facilitated the immense work which I have to do in order to give this dictionary the degree of exactness, clarity and usefulness which this new one should possess. When citing the degree of imperfection in the existing encyclopedia, I have mentioned in my prospectus only those matters pertaining to the form of the articles, their irregularity,their inappropriatenessfrom various points of view, etc., and, I assure you, I have in no way turned this picture into a caricature because I have remained silent about the essential insufficiencies of a great number of those articles which are filled with errors, misinterpretations,and wrong applications [of names]; my intention being not to criticize, in any of my publications, the authors who have written on subjects which I have to treat after them. It is not that I have not found numerous really interesting articles, because all those written by M. le Baron de Tschoudi* contain observations,even in great numbers, which I have used to advantage and which I shall not ignore. You will think, perhaps, that I have spoken too critically with respect to the dictionaries, but I beg you to take into account that natural history, and in particularbotany, does not lend itself with the same ease to a treatment in a dictionary as some other sciences, such as geometry, physics, chemistry, etc., especially if one takes care to mention in the main articles the various other articles which are relevant to them as well as the order in which they should be read, a point well stressed in your general scheme. For botany the great number of subjects is an insurmountable obstacle to the use of this device, because the number of cross-references and of articles to be cited would be enormous and such citations would still not dispel sufficiently the confusion caused by the alphabetical arrangement. This consideration has prompted me to abandon the most interesting aspects of my 'universal flora' in order to make the botanical dictionary, in the absence of a proper order, as useful as possible to the public. I am determined to give to this dictionary every value in my power for the benefit of the good that can come out of it and in acknowledgement of the trust that you have placed upon me. I have the honour, Monsieur, to be truly your most humble and obedient servant, le chevalier delamarck." * The Baron de Tschoudi, 'ancien Bailli de Robe-Court de Metz' wrote many of the articles on natural history, horticulture and arboriculturein the supplementaryvolumes of Diderot and d'Alembert'sEncyclopedie. Lamarck'scriticism is mainly directed against Daubenton and Adanson. AUGUST 1971 439 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions 2. Tableau encyclopddique et mdthodique des trois rdgnes de la nature. Botanique. Paris 1791-1823, 3 Tomes (6 volumes). Qu. The sixth column of the table of contents and dates of publication of this work given in my Taxonomic literature (1967), under no. 644, contains a serious typographical error; the correct information is as follows: 644. Tableau encyclopddiqueet mdthodiquedes trois regnes de la nature. Botanique. Paris 1791-1823, 3 Tomes (6 volumes). Qu. Publ.: Vols. 1-5(1) are by Lamarck, vol. 5(2) and tome 3 are by Poiret. The book is often cited as 'Illustration des genres' a title occurring on the introductory page of vol. 1, and the title-page of tome 3. tome I vol. 1 2 II 3 4 5 - III - part 1 2 1 2 1 2 1 2 1 2 - pages 1-200 201-352 353-440 441-496 1-40 41-48 49-72 73-136 137-551 1-728 plates 1-100 101-200 201-300 301-400 401-500 501-600 601-700 701-800 801-900 901-950 951-1000 - - 3 13 30 11 10 1 21 6 22 31 dates Mar 1791 Feb 1792 Jul 1792 Feb 1793 Apr 1794 Sep 1794 Nov 1796 Nov 1797 Dec 1799 Oct 1819 22 Feb 1823 22 Feb 1823 The plates are copper engravings of drawings by Audebert, Fossier, de Seve, Poiret fils and H.-J. Redout6. The originals were acquired by the Paris Museum d'Histoire naturelle in 1909. Ref.: BM 2: 528, NI 2244, PR 5005, RS p. 112, SK p. cxcv, SY p. 98 (index) IDC 445. Gleason, Phytologia 2: 292. 1947 (fasc. 60). Breistroffer,Proc. Verb. mens. Soc. Dauph. Ethn. Arch. 23 (nos. 182-184) Mar-Mai 1948. References and other relevant literature 1763 - Families des plantes, Paris. 2 vols. Facsimile reprint Lehre, with an introductionby Frans A. Stafleu. ADANSON, M. 1772 - Examen de la question: si les especes changent parmi les plantes; nouvelles experiences tentees A ce sujet. Histoire de l'Academie des Sciences 1769: 71-77, Memoires 1769: 31-48. AGASSIz,L. 1962 Essay on classification. Cambridge, Mass. (reprintof original edition of 1857). J. G. 1792 - Choix de memoires d'histoire naturelle. Paris. BRUGUi•m'REs, BUTLER, S. 1879 - Evolution, old and new; or the theories of Buffon, Dr. Erasmus Darwin, and Lamarck, as compared with that of Mr. Charles Darwin. London. CABANIS,P. J. G. 1802 - Rapportsdu physique et du moral de l'homme. Paris, 2 vols. A. P. DE 1862 - Memoires et souvenirs. Geneve. CANDOLLE, CAROZZI, A. V. 1964 - Hydrogeology by J. B. Lamarck [introductorynote and translation]. Urbana, Illinois. COLEMAN,W. 1964 - Georges Cuvier, zoologist. A study in the history of evolution theory. Cambridge, Mass. COSTANTIN, J. 1930 - Les derniers jours de Lamarck. Sa mort. Sa philosophie morale, Archives du Museum national d'Histoire naturelle s6r. 6.6: 5-10. G. 1807 - Eloge historique de Michel Adanson. Memoires de la Classe des CuvIER•, Sciences de l'Institut national de France 7(1): 159-188. 159. CUVIER, G. 1812 - Recherches sur les ossemens fossiles de quadrupedes. Paris, 4 vols. CUVIER, G. 1831 - Eloge de M. de Lamarck. M6moires de l'Academie royale des Sciences de l'Institut de France 13 (Hist.): i-xxxi. ADANSON, M. TAXON VOLUME 440 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions 20 DAUDIN, H. 1926 - Cuvier et Lamarck. Les classes zoologiques et l'idee de serie animale. Paris. DIDEROT,D. et J. D'ALEMBERT 1750-1780 - Encyclopedie, ou dictionnaire raisonn6 des sciences, des arts et m6tiers, par une soci6t6 de gens de lettres. Paris, 36 vols. Microfiche edition, Inter Documentation Company, Zug, no. 5437. GIARD, A. 1907 - Avant propos [to the reprint of Lamarck's Discours d'ouverture des cours de zoologie, in] Bulletin scientifique de la France et de la Belgique 40: 443-451. GILLISPIE, C. C. 1956 - The formation of Lamarck's evolution theory. Archives inter- nationales de l'Histoire des Sciences 9(4): 323-338. E. 1941 - Les sciences de la vie aux XVIIe et XVIIIe siecles. L'id6e d'6volution. Paris. HAMY, E. T. 1893 - Les derniers jours du Jardin du Roi et la fondation du Mus6um GUYENOT, d'Histoire naturelle. Paris. Les d6buts de Lamarck. Paris. G. 1905 - Un transformiste oubli6: Cabanis. Bulletin scientifique de la France HERV., et de la Belgique 39: 505-519. - Genera plantarum. Paris. Facsimile reprint Weinheim 1964, with JUssIEU, A. L. 1789 an introduction by F. A. Stafleu. KUEHNER, F. 1913 - Lamarck, die Lehre vom Leben. Jena. LACEPEDE, B. G. E. de 1800 - Histoire naturelle des poissons, tome second. Paris [pp. xxiii-lxiv, Discours sur la dur6e des especes]. *LAMARCK, J. B. A. P. M. DE 1778. Flore frangoise ou description succincte de toutes les plantes qui croissent naturellement en France. Paris, 3 vols. - Encyclopedie m6thodique. Botanique. Paris LAMARCK, J. B. A. P. M. DE 1783-1789 [vols. 1 and 2 are by Lamarck, vol. 3 is by Lamarck until the letter P for which he was assisted by Desrousseaux; the other volumes are by other authors]. LAMARCK, J. B. A. P. M. DE 1788 - M6moire sur les classes les plus convenables 'a 6tablir parmi les v6g6taux. Histoire de l'Acad6mie Royale des Sciences [Paris] 1785: 437-453. LAMARCK, J. B. A. P. M. DE 1790 - M6moire sur les cabinets d'histoire naturelle et HAMY, E. T. 1909 - particulierementsur celui du Jardin des plantes [Paris]. J. B. A. P. M. DE 1791-1799 - Tableau encyclop6dique et m6thodique des trois regnes de la nature. Botanique. Paris [vols. 1-5(1)]. Alternative title: Illustration des Genres. LAMARCK, J. B. A. P. M. DE 1792 - Journal d'histoire naturelle, r6dig6 par M. M. Lamarck, Olivier, Bruguieres, Haiiy, Pelletier. Paris, 2 vols. LAMARCK, J. B. A. P. M. DE 1794 - Recherches sur les causes des principaux faits physiques, et particulierement sur celles de la combustion. Paris, 2 vols. LAMARCK, J. B. A. P. M. DE 1796 - R6futation de la th6orie pneumatique. Paris. LAMARCK, J. B. A. P. M. DE 1797 - M6moires de physique et d'histoire naturelle. Paris. LAMARCK, J. B. A. P. M. DE 1798 - De l'influence de la lune sur l'atmosphere terrestre et les nuages. Journal de Physique 46: 428-435. - Annuaires LAMARCK, J. B. A. P. M. DE 1799-1810 m6t6orologiques. Paris, 11 vols. " Vers testac6s LAMARCK, J. B. A. P. M. DE 1798 - Tableau encyclop6dique ... coquilles bivalves. Paris. LAMARCK, J. B. A. P. M. DE 1800-1806 - Discours d'ouverture des Cours de Zoologie donn6s dans le Museum d'Histoire naturelle. An VIII, an X, an XI et 1806), [reprinted by A. Giard in] Bulletin scientifique de la France et de la Belgique 40: LAMARCK, 453-595. 1907. LAMARCK, J. B. A. P. M. DE 1801 - Systeme des animaux sans vertebres. Paris. LAMARCK, J. B. 1801 - Hydrog6ologie ou recherches sur l'influence qu'ont les eaux sur la surface du globe terrestre. Paris [see also Carozzi for English translation]. [N.B. published December 1801 or January 1802; title-page date 'An X.'] LAMARCK, J. B. A. P. M. DE 1802 - Recherches sur l'organisation des corps vivans. Paris (n.v.) * For full bibliographic details of Lamarck's publications see Landrieu, 1909. 441 AUGUST 1971 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions B. A. P. M. DE 1802-1806 - M6moires sur les fossiles des environs de Paris. [33 m6moires in the Annales du Museum d'histoire naturelle 1-8, in part reprinted in a single volume, 284 pp., Paris; for details see Landrieu 1909, pp. 460-461]. LAMARCK,J. B. A. P. M. DE 1809 - Philosophie zoologique, ou exposition des consid6rations relatives t l'histoire naturelle des animaux, a la diversit6 de leur organisationet des facult6s qu'ils en obtiennent. Paris. Facsimile reprint Weinheim 1960. LAMARCK,J. B. 1815-1822 - Histoire naturelle des animaux sans vertebres, Paris, 7 vols. Second edition, edited by G. P. Deshayes and H. Milne Edwards, Paris 1835-1845, 11 vols. LAMARCK, J. B. A. P. M. DE 1817-1818 - Conchiferes, Especes, Id6e, and other articles in Deterville, Nouveau dictionnaire d'histoire naturelle, Paris. 1816-1819, 36 vols. LAMARCK, J. B. 1820 - Systeme analytique des connaissances positives de l'homme, restreintes a celles qui proviennent directement ou indirectement de l'observation. Paris 1820 [re-issue 1830 with new title page]. LAMARCK, J. LAMARCK, J. B. et B. MIRBEL 1803 - Histoire naturelle des v6g6taux class6s par familles, vols. 1-2 [by Lamarck], Paris [unchanged reprint Paris 1825]. LANDRIEU, M. 1909 - Lamarck, le fondateur du transformisme,sa vie, son oeuvre. M6moires de la Soci6te Zoologique de France 21: 1-478. LECOMTE, H. et J. LEANDmRI1930 - L'Oeuvre botanique de Lamarck. Archives du Museum national d'Histoire Naturelle s6r. 6. 6: 31-44. LINNAEUS,C. 1751 - Philosophia botanica. Stockholm/Amsterdam.Facsimile reprint Lehre 1966. The great chain of being. Cambridge, Mass. [reprint Harper LOVEJOY,A. 1936Torchbook, New York 1960). MATOUT, M. 1930 - Lamarck m6t6orologiste.Archives du Museum national d'Histoire naturelle s6r. 6. 6: 45-48. J. P. 1963 - Adanson, the man. In G.H.L. Lawrence, editior, Adanson 1: NICOLAS, 1-122. Pittsburgh. A. S. 1901 - Lamarck, the founder of evolution. His life and work. New PACKARD, York. REVAULT D'ALLONNES, G. 1917 [?] - Lamarck, choix de textes et introduction. Paris (s.d.). J. 1963 - Les sciences de la vie dans la pensee frangaise du XVIIIme siecle. Paris. RcULE, L. 1930L'esprit Lamarckien dans les classifications actuelles. Archives du Museum national d'Histoire Naturelle s6r. 6. 6: 25-30. ROGER, SAINTE-BEUVE, C. A. 1834 - Volupt6. Paris. STAFLEU,F. A. 1963 - Adanson and the Familles des plantes. In G. H. M. Lawrence [ed.] Adanson 1: 123-264. Pittsburgh. STAFLEU, F. A. 1964 - Introduction to Jussieu's Genera plantarum. Weinheim 1964 (Historiae naturalis classica 35). STAFLEU, F. A. 1967 - Adanson, Labillardiere, de Candolle. Lehre. STAFLEU, F. A. 1967a - Taxonomic literature, A selective guide to botanical publications with dates, commentariesand types. Utrecht. F. A. 1971 - Linnaeus and the Linnaeans; the spreading of their ideas in systematic botany, 1735-1789. Utrecht (Regnum vegetabile vol. 69). STAFLEU, TREVIRANUS, G. R. 1802-1822 - Biologie oder Philosophie der lebenden Natur fiir Naturforscherund Arzte. Gottingen, 6 vols. TSCHULOK,S. 1937 - Lamarck. Eine kritisch-historische Studie. Ziirich und Leipzig. 442 TAXON VOLUME 20 This content downloaded on Mon, 24 Dec 2012 16:29:36 PM All use subject to JSTOR Terms and Conditions