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In his classic textbook, The History of Biology, Erik Nordenskiold suggested that ..... chapter on Humboldt's work in plant geography for Bruhn's commemorative.
British Society for the History of Science Humboldtian Plant Geography after Humboldt: The Link to Ecology Author(s): Malcolm Nicolson Source: The British Journal for the History of Science, Vol. 29, No. 3 (Sep., 1996), pp. 289-310 Published by: Cambridge University Press on behalf of British Society for the History of Science Stable URL: http://www.jstor.org/stable/4027735 Accessed: 16-12-2015 11:28 UTC REFERENCES Linked references are available on JSTOR for this article: http://www.jstor.org/stable/4027735?seq=1&cid=pdf-reference#references_tab_contents You may need to log in to JSTOR to access the linked references.

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BJHS, 1996, 29, 289-310

Humboldtianplant geographyafter Humboldt: the link to ecology MALCOLM

NICOLSON*

In his classic textbook, The History of Biology, Erik Nordenskiold suggested that there had existed, throughout the nineteenth century, not one but two distinct forms of plant geography.1 He designated one of these traditions of inquiry 'floristic' plant geography, tracing its origins back to the work of Carl Linnaeuson species and their distributions. The second form Nordenskiold termed 'morphological', by which he meant that its practitioners concentrated upon the study of vegetation rather than flora.2 He located the origins of this tradition of inquiry within the botanical work of Alexander von Humboldt. In a paper published in 1987, I presented evidence to support the thesis that Alexander von Humboldt's plant geography was indeed centrally concerned with vegetation - its character, distribution and relation to environmental parameters- and not solely or primarily with individual plants or species.3As Humboldt wrote in his Personal Narrative of Travels: I was passionatelydevotedto botany...and I flatteredmyselfthat our investigationsmightadd some new speciesto those whichhave beenalreadydescribed;but preferringthe connectionof factswhichhavebeenlong observedto the knowledgeof insulatedfacts,althoughtheybe new, the discoveryof an unknowngenusseemedto me far less interestingthanan observationon the geographicalrelationsof the vegetableworld,or the migrationof socialplants,and the limitof the heightwhichtheirdifferenttribesattainon the flanksof the Cordilleras.4 While a strong taxonomic component was necessarily retained within Humboldt's botanical practice, it was toward the elucidation of these supra-specific matters that his botanical investigations were principally directed. There was thus, as Nordenskiold had outlined, a striking difference between Humboldt's principal objectives and those of the Linnaean botanists. *

Wellcome Unit for the History of Medicine, 5 University Gardens, Glasgow G12 8QQ. 1 E. Nordenski6ld, The History of Biology, New York, 1928, 560-1. 2 The flora of a given region consists of the plant species which grow there. Students of floristics may make generalizations about the characterof a region's flora as a whole, but the level of analysis on which their scientific practice is based is essentially that of the individual species or genera. Vegetation is a collective phenomenon produced by many species together. The characteristicsof vegetation are produced not only by the presence or absence of particular species, but also by their different growth forms and relative abundances. For a precise elucidation of the distinction between the study of flora and vegetation, see F. E. Egler, 'Vegetation as an object of study', Philosophy of Science (1942), 9, 245-60. 3 M. Nicolson, 'Alexander von Humboldt, Humboldtian science, and the origins of the study of vegetation', History of Science (1987), 25, 169-94; see also note 5 below. 4 A. von Humboldt, Personal Narrative of Travels to the Equinoctial Regions of the New Continent 1799-1804 (tr. H. M. Williams), 6 vols., London, 1821-25, i, p. iii.

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Humboldt's plant geography was distinctively vegetational geography. Not only did vegetation constitute for Humboldt an object of inquiry per se, it was also conceived of as existing in regional or topographical types or units. This assumption that there were natural kinds of vegetation was one of the key elements in the rationale of the new practice. To Humboldt, moreover, plant geography was not merely a major branch of natural history. It was a crucial link between the natural sciences and the human sciences. It was a central element within Humboldt's programmefor 'la physique gene'rale'- the universal science which would encompass all natural phenomena, and aesthetics and epistemology besides.5 This paper aims to establish that this type of vegetational plant geography was not merely a personal idiosyncrasy of Alexander von Humboldt but that it constituted a distinctive tradition of inquiry, which persisted, developed and diversifiedthroughout the nineteenth century. Humboldtian plant geographers are identified in Scandinavia, the German-speakingcountries and France.6It is also argued that Humboldtian plant ecology provided one of the principal elements from which a newly 'self-conscious' science of ecology was composed, at the end of the century.7 ALEXANDER

VON HUMBOLDT'S

PLANT GEOGRAPHY

Humboldt first publicly set out his programmefor a new form of plant geography in 1793. In his Flora Fribergensis specimen, he followed Kant in distinguishing between a true history of nature and an artificial description of nature such as had been provided by the older, Linnaean system of natural history.8 No longer should botanists study merely individual species and their outward appearances; no longer should they be preoccupied solely with descriptive taxonomy and nomenclature. The central concern of the botanist, or rather the plant geographer, should be, by contrast, the collective, holistic phenomena of vegetation: Observationof individualpartsof treesor grassis by no meansto be consideredplantgeography; ratherplantgeographytracestheconnectionsandrelationsbywhichall plantsareboundtogether amongthemselves,designatesin whatlandstheyarefound,in whatatmosphericconditionsthey live... and describesthe surfaceof the earthin whichhumusis prepared.9 5 See M. Nicolson, 'Alexander von Humboldt and the geography of vegetation', in Romanticism and the Sciences (ed. N. Jardine and A. Cunningham), Cambridge, 1990, 169-88; also M. Nicolson, 'Historical introduction', in Alexander von Humboldt, Personal Narrative of Travels (tr. J. Wilson), London, 1995; and M. J.Bowen, Empiricism and Geographical Thought: From Francis Bacon to Alexander von Humboldt, Cambridge, 1981. 6 I have, unfortunately, been unable to investigate the presence or otherwise of Humboldtian traditions in other areas where distinctive schools of plant sociology developed, for example the Netherlands and, notably, Eastern Europe. 7 The term 'self-conscious' to distinguish the science of ecology, so-called, from pre-existing knowledge and investigations of the environmental relations and interactions of plants and animals, is taken from W. C. Allee, A. Emerson, T. Park, 0. Park and K. Schmidt, Principlesof Animal Ecology, Philadelphia, 1949, 1-59. For other discussions of the place of Humboldt's plant geography within the development of ecology, see P. Acot, Histoire de l'e6cologie,Paris, 1988, and J. Drouin, Re'inventerla nature: L'e'cologieet son histoire, Paris, 1991. 8 See J. A. May, Kant's Concept of Geography, Toronto, 1970. 9 A. von Humboldt, Flora Fribergensisspecimen, Berlin, 1793, 9-10. The translation is from R. Hartshorne, 'The concept of geography as a science of space, from Kant and Humboldt to Hettner', Annals of the Association of American Geographers (1958), 48, 97-108, on 100.

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This grandvisionof a completehistoricalgeographyof the earthprovideda centraltheme for Humboldt'slater work. The importanceof geographywithin Humboldt'sinvestigativeenterprisewas also expressedin thepriorityhe accordedto scientifictravelling.To Humboldtexplorationwas an essentialpartof naturalinquiry,a necessaryconditionof 'la physiquege'ne'rale'. Of all his travelsHumboldtregardedthe journeyto the Americasas the most importantsince, in the New World,he had visitedthe Tropicswhereplant and animallife displayedthe greatestvarietyand richness.Withinhis accountsof his travelsin SouthAmerica,Mount Chimborazooccupiesa placeof specialsymbolicimportance.In 1802,Humboldtand his companion,AimeBonpland,attemptedto climbthe mountain,ascendingfromthe levelof rainforestand humansettlement,throughthe severalaltitudinalvegetationzones, to the region of permanentsnow, taking measurementsall the way. On its slopes they thus experienced,withina smallcompass,muchof the physicalandvegetationaldiversityof the continent.In theircampat the foot of the mountainHumboldtbeganto composea fuller articulationof the programmefor plant geographyhe had adumbratedin the Flora Fribergensisspecimen.The Essaisur la ge'ographie des planteswas publishedin 1807.10 Humboldt's plant geography was a thoroughly empirical investigation of the environmentof plants. Instrumentswere used intensivelyto measurea wide varietyof physicalparameters.'1 The readingsweretabulated,comparedbetweendifferentsites,and correlatedwith the occurrenceof the varioustypesof vegetation.It was hopedthat such correlationswould aid in the discernmentof the laws whichgovernedthe distributionof vegetation,revealingthe distinctiveintegrativeprocesseswhich formed 'the natural divisionsof the vegetableempire'.12 His concernwith the unity of landscapeis well exernplifiedby the 'Tableau physiquedes Andes et pays voisins'. This is a large and elaborateengraving,foldedwithinthe pagesof the Essai,whichdepictsa cross-sectional profileof the Andesfromthe Atlanticto the Pacificat the latitudeof Chimborazo.In this one figurearemappedor tabulated:whichplantandanimalspecieslive where,wherethe altitudinalzones of vegetationbegin and end, the types of agriculturepursued,the underlyinggeologicalstructures,and a wide varietyof physicalor meteorologicaldata. The objectwas to give, in a singleillustration,a completeimpressionof a naturalregion - the 're'gionse'quinoctiales' of SouthAmerica.The 'Tableau',with its holisticvision of a complexyet unifiedlandscape,is a verytypicalHumboldtianproduction. The 'Tableauphysiquedes Andeset pays voisins' also pictoriallyrepresentedspatial differentiation withina singleregion.Withinthe 're'gionsequinoctiales'werefound,high on the mountains,a 're'giondes lichens'and,lowerdown,a 're'giondes Cinchona'.These smallersortsof vegetationalregionweredistinguishable by physiognomiclife-form,by the generalappearanceand habit of growthof the constituentplants.'Re'gionsdes lichens' were distinguishableby the obviousprofusionof a numberof species,all with the same lichenous life-form. The study of plant physiognomywas an importantfeature of Humboldt's botanical enterpriseand this aspect of his investigationof vegetation 10 A. von Humboldt, Essai sur la ge'ographiedes plantes, Paris, 1807. 11 For the instrumental nature of Humboldtian science, see S. F. Cannon, 'Humboldtian science', in Science in Culture: The Early Victorian Period, New York, 1978, 73-110. 12 Humboldt, op. cit. (4), i, 158.

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constitutes one of the most decisive ways in which he departed from Linnaean taxonomic methods. Classificationby life-form, although in some cases it might approximate to more orthodox arrangements,was essentially independent of floristic systems: In determiningthose forms, on whose individualbeauty, distributionand grouping, the physiognomyof a country'svegetationdepends,we must not groundour opinion...on the smallerorgansof propagation...but mustbe guidedsolelyby thoseelementsof magnitudeand mass from whichthe total impressionof a districtreceivesits characterof individuality...The systematisingbotanist...separatesinto differentgroupsmanyplantswhich the studentof the physiognomyof natureis compelledto associatetogether.'3 Similarly,species closely allied by the taxonomist might be very differentphysiognomically. One of Humboldt's reasons for according a central importance to plant geography within the scheme for a universal science was his conviction that the vegetation of any given region was not only a primary expression of the physical environment; it also exercised a formative influence on humanity, both materially and spiritually.'4 The vegetation was not, of course, the only feature of the environment which was morally or aesthetically influential. But the role played by plants en masse in mediating between mankind and the physical environment was a major one: Howevermuchthe characterof differentregionsof the earthmaydependupon a combination of all theseexternalphenomena... the outlineof mountainsandhills,the physiognomyof plants andanimals,the azureof the sky,theformsof thecloudsandthetransparency of the atmosphere, still it cannot be deniedthat it is the vegetablecoveringof the earth'ssurfacewhich chiefly conducesto the effect.'5 A HUMBOLDTIAN TRADITION - SCHOUW, MEYEN, GRISEBACH AND KERNER VON MARILAUN Humboldt never held an academic position so he had no students as such. However, his voluminous published work provided vivid and detailed exemplifications of the new form of botanical inquiry. He exercised major influence through a vast scholarly acquaintance and he had at his disposal considerable academic patronage, both in Paris and Berlin."6He maintained an immense scientific correspondence.'7One of his regularcorrespondents,the Dane Joachim F. Schouw, may be identified as one of the earliest Humboldtian plant geographers. Schouw read and was inspired by Humboldt's work in the early 1810s, when he was abandoning a career in law and training to become a botanist.'8 He was applying Humboldtian principles to the investigation of the vegetation of Scandinavia by the end of the decade. This is indicated by the title of one of his first publications, 'Einige 13 A. von Humboldt, 'Ideas for a physiognomy of plants', in Views of Nature or Contemplations on the Sublime Phenomena of Creation (tr. E. C. Otto and H. G. Bohn), London, 1850, 220-1. 14 Humboldt, op. cit. (10), 30-1. All translations from this work are my own. 15 Humboldt, op. cit. (13), 217. 16 See E. R. Braun, Alexander von Humboldt: Patron of Science, Madison, 1954; and I. Jahn, 'The influence of Alexander von Humboldt on young biologists and biological thinking during the 19th century', Actes du XIe Congres International d'Histoire des Sciences, Warsaw, 6 vols., Warsaw, 1965, v, 81-6. 17 See H. Beck, Gesprache Alexander von Humboldt, Berlin, 1959. 18 A. P. M. Sanders, 'Schouw, Joachim Frederik', DSB, xii, 214-15.

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betreffendeWerke des Herrn von Bemerkungenfiber zwei, die Pflanzengeographie In 1819Schouwtravelledto Paristo meethis mentoron the way backfrom Humboldt'."9 a scientificjourneyto Italy.He travelledin Italyagainin 1829,laterpublishingan account of theseexpeditions,Tableaudu climatet de la vegetationde l'Italie.This did not appear until 1839. In his introductionto the work he apologizedfor his delayin producingan account of his Italian investigations.This, he said, was due to his absorptionwith Humboldtianconcernsin his own country,with 'son traitessur la geographieuniverselle des planteset son tableaudu climatdu Danemark'.20 Schouw'sdescriptionsof his own workillustratehow similarhis concernswereto those of Humboldt.Schouwcarefullymeasuredthe physicalenvironmentto correlateit withthe distributionof vegetation: Le volume actual [the first]contientle tableaude la temperatureet des pluies, et commela du sol exerceune influenceessentiellesurle climat... Le secondvolumecontiendra configuration les autreselemensdu climatet une comparaisondes anneesdiversesrelativementau caractere m'teorologique,ce qui conduiraaussi a traiterla question interessantedes variationsqu'a attribueau climat.Le troisiemevolumeseraconsacreau tableauphytogeographique.

The firstvolumewas given over entirelyto tablesof physicalmeasurement: pas l'elevationdu sol au-dessusde la mer, Le climatet la vegetationsont influencespuissamment de la hauteurd'ungrandnombre le but de mon voyageexigeoitparconsequentla determination de points.22

As faras I havebeenableto discern,Humboldtwas thefirstto callcommunitiesof plants ' associations'.Humboldtusedthe terminformally- 'The heaths,thatassociationof erica 23 But, among vulgaris,of ericatetralixand the lichensicmadophiliaand haemotomma'. Humboldtianplantgeographers,the term'association'becamea technicalone meaninga Schouwwas particularly more or less definiteand distinguishableplant community.24 interestedin the unitsof vegetation.In 1822,he inventeda nomenclaturefor associations which consistedof adding the suffix '-etum' to the genericname of the plant which dominated the given association. For example, 'Fagetum' indicated a community dominatedby a speciesof Fagus,the beechtree;'Quercetum',an associationin whichone of the speciesof the oak, Quercus,was the most numeroustree.25 unit of The Humboldtianprogrammefor a plantgeographybasedon a supra-specific botanists, vegetationwas also continuedand developedin the work of German-speaking notablyFranzMeyen,AugustGrisebachand Kernervon Marilaun.In theirwork we can 19 J. F. Schouw, 'Einige Bemerkungen uber zwei, die Pflanzengeographiebetreffende Werke des Herrn von Humboldt', Jahrbucherder Gewacbskunde (1818), 1, 6-56. 20 J. F. Schouw, Tableau du climat et de la vig6tation de l'Italie, Copenhagen, 1839, p. viii. I have been unable to trace the second and third volumes of this work. 21 Schouw, op. cit. (20). 22 Schouw, op. cit. (20), p. ix. 23 Humboldt, op. cit. (10), 17. 24 This argument is set out in more detail in M. Nicolson, 'The Development of Plant Ecology, 1790-1960', Ph.D. thesis, University of Edinburgh, 1984, 50-1. 25 J. F. Schouw, Grundzuge einer allgemeinen Pflanzengeograpbie,Berlin, 1823, 165. The nomenclature was apparentlyfirst proposed in an earlier (1822) Danish edition of this work, see R. H. Whittaker, 'Classification of natural communities', Botanical Gazette (1962), 28, 1-239, on 9.

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see the full impact of the Humboldtian legacy as it was expressed in the scientific practice of the following generation of research workers. Franz J. F. Meyen was, until his death in 1840, Professor of Botany at the University of Berlin. He was one of the most favoured of Humboldt's many scientificproteges.2"Meyen, who had been working as a physician when Humboldt returned to Berlin in 1826, owed both his scientific career and his post at the University to Humboldt. Humboldt also took great trouble to make Meyen's work known to as wide an audience as possible, personally sponsoring the Frenchtranslation of Meyen's book on plant geography.27In return,Meyen regularly invoked Humboldt's name throughout his writings. Meyen's Grundriss der Pflanzengeographie represents one of the earliest, most successful and most explicit articulations of the Humboldtian exemplar. Meyen presented his plant geography as 'the new science which answers in a way that had before been impossible, many of the most interesting questions on the production and distribution of organic beings on the surface of the globe'.28 In some respects, Meyen's career was almost a replica in miniature of Humboldt's. Meyen, for example, travelled in the New World between 1830 and 1832. Like Humboldt, he recorded both scientific and aesthetic observations on the summits of the Andes: 'The sight of a little Gentian, very similar to our Gentiana uliginosa and G. nivalis at the height of 14,000 or 15,000 feet as in the Cordilleraof SouthernPeru, can enchain the botanist for hours; he again and again gathers this little plant which takes him, at least in imagination, home.'29 Extant correspondencebetween Humboldt and Meyen indicates that Humboldt advised his protege upon his travel plans at every stage. Outlines of the Geographyof Plants, to give Meyen's book its English title, was, among other things, a sustained and sophisticated attempt to correlate vegetation with measured physical factors. Meyen employed copious instrumental records, mostly of his own making, and drew up detailed temperature charts and tables. He employed the isoline mapping method, as developed by Humboldt, to apply meteorological data to the study of the distribution of vegetation:30 It is veryeasyto showthatthe conditionsof climate,particularly heatandmoisture,arethe chief causeswhichdeterminethe stationand distributionof plantsand thereforeit is of the greatest importanceto knowexactlythe modesin whichthe influenceof the oftenextremelycomplicated conditionsof climatebecomeapparent.To arriveat this end, we mustfirst... employourselves with the observationswhichhavebeencollectedon the distributionof the heat and moistureof the atmosphereoverthe wholeglobeandwhichareby no meansof puremeteorologicalinterest but constantlypoint to the influencewhichindividualmeteorologicalphenomenaexerciseover vegetation.31

Meyen was interestedin floristics and systematics, but he carefully distinguishedfloristic plant geography from vegetational plant geography, which interested him still more. 26 See Jahn, op. cit. (16). 27 F. J. F. Meyen's text was translated into English as Outlines of the Geography of Plants (tr. M. Johnston), London, 1846. 28 Meyen, op. cit. (27), 3. 29 Meyen, op. cit. (27), 27. 30 See A. H. Robinson and H. M. Wallis, 'Humboldt's map of isothermal lines: a milestone in thematic cartography', CartographicJournal (1967), 5, 119-23. 31 Meyen, op. cit. (27), 8.

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Withinthe studyof vegetation,the classificationof physiognomywas one of his principal concerns: The subjectof the distributionof plantsover the surfaceof the globe may be dividedinto two considersvegetationaccording perfectlydistinctbranches,one of whichcalledthePhysiognomics to the distributionof formswhichpointout the groupsof plants;it is a peculiarlynaturalsystem in whichsimilarityof form is the principleof classification.32

ExplicitlyfollowingHumboldt'sexample,Meyenemphasizedtheroleplayedby vegetation withinMan's aestheticappreciationof Nature: ... celebratedwork 'Considerationson the Physiognomyof BaronAlexandervon Humboldt['s] Plants'pointedout thishighlyinterestingsidefromwhichbotanymaybe viewedandhow it may improvethe tasteof nationsby increasingtheirsensibilityto the beautiesof natureandthushave an influenceon the progressof the arts.33

Meyen also consideredthe effect of vegetationupon human society, developinghis mentor'sargumentthat the regionalityof vegetationwas both a reflectionand a causeof the naturaldivisionsof geographicalphenomena: It is vegetationwhich fixes the naturalcharacterof a region and determinesthe conditions accordingto which men gatherinto varioussocieties,at one time leadinga nomadiclife, at Wherevegetationis scanty, anotherenjoyingmoreor less the beneficentinfluenceof agriculture. andmanis moreor less confinedto animalfood, as in the caseof the SamoyedesandEsquimaux on the coastof the Northernocean,civilisationis impossible.In thoseregions,manliveslikethe beastsand does not even thinkof rousinghimselfabovethem.34

AugustGrisebachwas bornin 1814.35His earlyinterestin botanywas encouragedby his uncle, the eminentnaturalhistorianGeorgF. W. Meyer,who was a professorat the Universityof Gottingen,whereHumboldthadstudied.Grisebachwas himselfa studentat Gottingenfrom1832until1834whenhe movedto the Universityof Berlin,studyingunder FranzMeyen.He also becameacquaintedwith C. S. Kunth,who had been the botanist principallyin chargeof the taxonomicwork upon the specimenscollectedby Humboldt and Bonplandin SouthAmerica.36 Althoughprincipallya taxonomist,Kunthwas, thus, one of Humboldt'sclosestscientificcollaborators.Grisebachbecame,like Meyen,a close associateof Humboldtandhe too self-consciouslycontinuedHumboldt'sprogrammefor a vegetationalplant geography. In 1833Grisebachwent on a botanizingtrip to southernFrance.He had alreadyread Humboldt'saccountof his SouthAmericanexpeditionand was interestedin the typesof of differentregionsanddifferentenvironments.In 1839he set out vegetationcharacteristic regionsof Thrace, againon a majorexpeditioninto the hithertobotanically-unexplored 32 Meyen, op. cit. (27), 98. 33 Meyen, op. cit. (27), 98. 34 Meyen, op. cit. (27), 1. 35 The best biographical source for Grisebach in English is W. T. Stearn, 'Grisebach's Flora of the British West Indian Islands: a biographical and bibliographicalintroduction', Journal of the Arnold Arboretum (1965), 46, 243-85; see also T. A. C. Balfour, 'August Heinrich Rudolph Grisebach', Transactions of the Botanical Society of Edinburgh (1882), 14, 13-20. 36 For Kunth and his association with Humboldt, see W. T. Stearn, 'Carl Sigismund Kunth', in Humboldt, Bonpland, Kunth and Tropical American Botany (ed. W. T. Steam), Lehre, 1968.

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Macedonia, Albania and Northern Asia Minor. He made a transect across the Balkan Peninsula, very similar to the transect of South America pictorially represented in Humboldt's Essai. These travels were described in his Reise durch Rumelien und nach Brussaim Jahre 1839.37This account contains much Humboldtian observation,particularly of altitudinal zonation of vegetation. For example, on the slopes of Ulu Da', he described three main zones of vegetation - the region of sweet chestnuts, the region of conifers, and the alpine region. On his return to Germany Grisebach became privatdozent at the University of Gottingen, rising to full professor in 1847. Another botanizing and phytogeographical journey was undertaken, to Norway, in 1842. Grisebach became a personal friend of Humboldt and the two maintained a copious scientific correspondence. As noted above, Schouw had devised a system of nomenclature for vegetational entities based upon their floristic characteristics.In 1838, however, Grisebach re-emphasized the physiognomic aspect and introduced the new vegetational term 'formation': I wouldterma groupof plantswhichbearsa definitephysiognomiccharacter,suchas meadow, by a singlesocial formation.The lattermay be characterized a forest,etc., a phytogeographic species,by a complexof dominantspeciesbelongingto one family,or finally,it may show an aggregateof species, which, although of various taxonomic character,have a common peculiarity;thus the alpinemeadowconsistsalmostexclusivelyof perennialherbs.38 It became conventional to use the term 'association', often together with the '-etum' suffix, to refer to vegetation types characterizedby floristic criteria, and the term 'formation' to refer to types characterized by physiognomy, as in Grisebach's examples. Grisebach was an accomplished taxonomist and he was interested in floristic as well as vegetational plant geography. Indeed the two activities often complemented each other: 'Bunbury['s] description of the characterof the vegetation in the environs of Cape Town is so much the more interesting, as being accompanied by Harvey, who is intimately acquainted with the Cape Flora, he was enabled to acquire an exact knowledge of the species'.3 But Grisebach'sapproach to plant geography was characterizedby a distinctive emphasis on vegetational phenomena and their relation to the physical environment. A good example of this is afforded by his description of the plant cover of European Russia: NorthernRussiais chieflydistinguishedfromthe centralprovinceby its denseforests,in which Pinussylvestris.L., and P. abies, L., arethe predominantspecies,and whosevast extentis only brokenby swamps... The pines and firs form two distinctforest formations,differingin the proportionof the argillaceousconstituentof the soil. The clayey,often marshylow landsof the old red sandstoneare coveredwith thickfirwood, amongwhichoccurthe aspenand the alder; the sandydiluvialhillocksbearPinussylvestris,L., and Betulapubescens,Ehrb.,and represent the forestcharacterof the North Germanplain,the soil of whichhas beenformedat the same time. On this diluvium,wherethe soil is deficientin clay, are met with also heathsof Calluna whichdo not occurin the Siluriaplainsand trapformations.40 37 A. H. R. Grisebach, Riese durch Rumeliaen und nach Brussa im Jahre 1839, 2 vols., Gottingen, 1843-46. 38 A. H. R. Grisebach, 'Ueber den Einfluss des Climas auf die Begranzung der naturlichen Floren', Linnaea (1838), 12, 159-200, on 160. The translation is from F. E. Clements, Plant Succession: An Analysis of the Development of Vegetation, Washington, DC, 1916, 116-17. 39 A. H. R. Grisebach, 'Report on the contributionto botanical geography during the year 1842', Reports and Papers on Botany, Ray Society (1846), 57-122, on 96. 40 A. H. R. Grisebach, 'Report on the contributionto botanical geography during the year 1843', Reports and Papers on Botany, Ray Society (1846), 125-212, on 132-3.

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Whilefarthersouth: The northernmarshwillows are replacedby Salix fusca, L., Cinerea by Caprea L., and Alnus incona, D.C., is represented by Alnus glutinosa, G. Thusalmostall the plantformationsassume anothercharacter,but the physiognomyof the wholecountryis muchmorestrikinglyalteredby the increasedextentof cultivation.41

Grisebach'sprogrammeof researchin Humboldtianplantgeographyculminatedwith the publicationin 1872of the monumentalDie Vegetationder Erde,the firstattemptto A total providea comprehensive descriptionandclassificationof the world'svegetation.42 of fifty-fourphysiognomicgroupswererecognized.Grisebachmaythereforebe confidently identifiedas one of the leadingexponentsof Humboldtianplantgeography.His statusas one of Humboldt'sleadingbotanicaldiscipleswas acknowledgedby his being askedto write the chapteron Humboldt'swork in plant geographyfor Bruhn'scommemorative scientificbiography.43 In his classictext Das Pflanzenleben derDonauldnder,publishedin 1863,AntonKerner von Marilaun,Professorof Botanyat the Universityof Innsbruck,notedthatowingto the scientifictravelsof Humboldtmorehad beenknown of the vegetationof SouthAmerica than of Austria-Hungary : Attentionwas calledto our nativeplantformationsonly whentravellerswith geniusand good fortuneshowedin word and picturethe marvellousplantformationswhichspreadin primeval virginitybeneathtropicsunsalongthe banksof the giantriversof SouthAmerica...It is literally truethatwe havehadexactdescriptionsandsplendidlyillustratedportrayalsof the shoresof the PacificOceanor thetropicalzoneof Brazilfora longtimebeforeournativeplantformationswere given a similartreatment.45

taskwas to redressthisimbalance,usingHumboldt's Kernervon Marilaun'sself-appointed own methods.He travelledextensivelyin the previouslybotanicallyunexploredregionsof easternHungaryand Transylvaniato studythe types of vegetation: It mustbe one of the most interestingand importantservicesof Botanyto devotemoreprecise observationthanhas hithertobeengivento the lawsgoverningthe structureanddevelopmentof plantformationsand theirrelationto local climaticconditions.It is the plantformationswhich lineament:it is they whichproducea livelyimpression give to everylandscapeits characteristic by reasonof greatvariabilityandrapidlychangingcontrastsor givea senseof monotonybecause of few and simpleforms.46

In the characterizationof vegetationthe criteriahe used were physiognomicand he proposeda new nomenclaturefor plant formationsbasedon dominantlife-forms.Like Meyen,he was carefulto distinguishhis preferredstudyof physiognomyfromthe study of floristics:'almosteverysystemicgroupis represented by variousformswhicharetotally 41 Grisebach, op. cit. (40), 135. 42 A. H. R. Grisebach, Die Vegetation der Erde nach ihrer klimatischen Anordnung, Leipzig, 1872. 43 A. H. R. Grisebach, 'Pflanzengeographie und Botanik', in A. von Humboldt: Eine wissenschaftliche Biographie (ed. K. Bruhns), 3 vols., Leipzig, 1872, iii, 233-68. 44 For biographical details of Kernervon Marilaun, see E. M. Kronfeld, Anton Kernervon Marilaun: Leben und Arbeit, Leipzig, 1908. 45 A. Kernervon Marilaun, Das Pflanzenlebender Donauldnder,Innsbruck,1863. I have used H. C. Conard's translation, The Backgroundof Plant Ecology: A Translationfrom the Germanof' The Plant Life of the Danube Basin', Ames, 1951, 5. 46 Von Marilaun, op. cit. (45), 10.

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different in physiognomy ... Plant physiognomy and plant systematics go entirely different ways'." To Kernervon Marilaun, as to Grisebach, the individual plant was fully understandable only if it was considered as a member of a distinct community. As well as being a unified response to the climate and other environmental factors, the plant community displayed a high degree of internal integration. This was part of the reason one community was readily distinguishable from its neighbours. Plants living together had effects upon one another and the characterof vegetation was partially determinedby these interactions. The social groupings of plants had definite structure: is by no meansaccidentalin spite Thehorizontalandverticalassortingof largeplantcommunities of its apparentlackof order.It followscertainimmutablelaws.Everyplanthasits place,its time, its functionand its meaning...In everyzone the plantsare gatheredinto definitegroupswhich appeareitheras developingor as finishedcommunities,but nevertransgressthe orderlyand correctcompositionof theirkind.Sciencehas givento suchgroupsthe namePlantFormations. Withthe comparativestudyof landscapesbotanistsfoundit necessaryto defineandcharacterize elementswhichare so conspicuousin the physiognomyof landscape.48 theseever-recurring Following Humboldt, von Marilaun was concerned to interpret vegetation as an expression of the physical environment: 'Everywhere plant life is adjusted to the local climatic conditions. When one opens the great green Book of Nature, one finds therein the local climatic conditions generally much more precisely and correctly registered than on the yellowing pages of thick meteorological journals and folios. The vegetation is everywhere the reflection of the local climate. For that reason no feature of the landscape is so significant and informing as the vegetation'.49 He emphasized, moreover, that vegetation affected the entire range of human activities: 'What a wealth of problems arise in investigating the relation of these differentexpressions of landscape to the spiritual side of Man, to his feeling toward nature, to his culture and to the products of his creative art' 50

It is clear from Das Pflanzenlebender Donauldnder that, by the middle of the nineteenth century, much more was known about vegetation than when Humboldt wrote his Essai. Marilaun's descriptions of the plant formations of Austria-Hungary were very detailed his accounts of plant succession, what he termed 'the genetical relationships of plant formations', were particularlywell-observed.5' He could also confidently point to several instances where the study of vegetation had proved its practical utility to fields such as forestry. But the framework of his inquiry remains recognizably Humboldtian. SOUTHERN

HUMBOLDTIANS

- LECOQ AND HEER

In France and especially in Britain, an alternative floristic style of botanical geography seems to have been the most popular. Watson, Hooker and Wallace, to name but three, all practised a form of botanical investigation principally structuredaround the problems 47 48 49 50 51

Von Von Von Von Von

Marilaun, op. Marilaun, op. Marilaun, op. Marilaun, op. Marilaun, op.

cit. cit. cit. cit. cit.

(45), p. v. (45), 4-5. (45), 3. (45), 10. (45), 196-205.

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of distributionand dispersalof species.62There appearto have been few nineteenthwho placedgreatemphasison vegetationalphenomena.63 centuryBritishplantgeographers countriesandin Scandinavia,the Humboldtianstyleof However,in the German-speaking vegetationalplant geographyseems to have been widely adopted.It is instructive,for text with the mostHumboldtiansoundingtitle, example,to comparethe English-language TheRegionsof Vegetation:Beingan Analysisof theDistributionof VegetableFormsover the Surfaceof the Globe,RichardHind'saccountof his botanicalobservationsduringthe voyageof HMSSulphur,whichwas publishedin 1843,with the roughlycontemporaneous work of OswaldHeer, Professorof Botanyat the Universityof Zurich.54 Despitehavingentitledhis book The Regionsof VegetationHind basedhis discussion upon what are somewhatarbitrarygeographicaldivisions- the Japanregion,the China region,the Birmahregionandso forth.56His descriptionsof theseregions,apartfrombrief remarksas to the generalcharacterof the landscape,were entirelyfloristic.Only in his accountof the 'Himma-lehregion'did he departfromthis procedure.Here,with a bow, as it were, to Humboldt'sfamousdescriptionof the Andes,he recognizedthe 'regionof lowlandcultivation', the 'regionof woods', the ' regionof shrubs', the 'regionof grasses' and the 'region of cryptogamicplants'. The emphasisof Oswald Heer's Beitragezur on the otherhand,was almostentirelyvegetational.Foldedin at the Pflanzengeographie, book is an endof the engravingof a mountainpanorama,entitled' GemdldederVegetation des SudostlichenTheils des CantonGlarus'.This is strikinglysimilar,in size, style and the content,to Humboldt's'Tableauphysiquedes Andeset paysvoisins'.Accompanying engravingis an elaboratetableindicatingthephysicalenvironmentof eachregiondepicted. for instance,are given.Symbolson the soil and atmospherictemperatures, Characteristic pictureindicatethe most importantspeciesto be foundin each region.For instance,the rocksof the upperzone of the 'regionivalis'aresaidto supporta vegetationcharacterized by saxifragesand Primula. Heerwas carefulto associatehimselfwith the Humboldtiantraditionandto distinguish his approachto plantgeographyfromthat of the morefloristicLinnaeanbotanistGeorg von denPflanzenaller Wahlenberg:'Ich suchteaufdiesenReisenmirgenaueVerzeichnisse wobeiich die Methodevon Schouwbefolgte... Die H6henundLocalitatenzu verschaffen,

52 There is, however, a short description of the vegetation of the British Isles, more or less along the lines of the Continental model, within the floristic text: H. C. Watson, Remarks on the GeographicalDistribution of the British Plants, London, 1835, 34-60. 53 It is instructive that, for example, Janet Browne, in her history of biogeography, concerns herself only with studies of the distribution of species and mentions no Continental biogeographer after Alphonse de Candolle; J. Browne, The Secular Ark: Studies in the History of Biogeography, New Haven, 1983. The insularity of British biogeography after 1840 has been noted by G. Nelson, 'From Candolle to Croizat: comments on the history of biogeography', Journal of the History of Biology (1978), 9, 269-305. Strangely, however, Nelson is unable to specify precisely what British biogeography was isolated from. It may be worth investigating whether it was isolated from a distinct Continental tradition of Humboldtian biogeography. See also J. Browne, 'A science of empire: British biogeography before Darwin', Revue d'Histoire des Sciences (1992), 45, 453-75. 54 For biographical details of Heer, see Anon., 'Obituary - Dr Oswald Heer', Proceedings of the Linnean Society of London (1883-86), 84. 55 R. B. Hinds, The Regions of Vegetation: Being an Analysis of the Distribution of VegetableForms over the Surface of the Globe, London, 1843.

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Methode Wahlenberg fand ich sehr unzweckmassig'.` That Heer should distinguish his work from that of Wahlenbergis instructive.The holder of the Linnaean chair in Uppsala, Wahlenbergwas not merely a collector and taxonomist. Unlike the majority of nineteenthcentury Linnaean botanists, Wahlenberg sought to develop the form of inquiry into the interrelationsbetween animals and plants in their natural environment which had been set out by Linneaus in such works as Oeconomia naturae and Politae naturae.57Wahlenberg investigated the distribution of plant species in relation to a number of environmental variables, including underlyinggeology and mineralogy, climate, and soil composition and temperature.With the benefit of hindsight, one may discern in Wahlenberg'sresearchesthe essentials of a programme of autecological plant geography. But Wahlenberg established no lasting school or tradition. Those botanists who wished to study the relations of plant species to each other and to the physical environment generally preferred, like Oswald Heer, to articulate the Humboldtian rather than the Linnaean exemplar. In 1849, the Swiss botanist Jules Thurmann made explicit the distinction between the study of floristics and the study of vegetation: A region'sfloracomprisesthe enumerationanddescriptionof all its specieswithoutreferenceto theirabundance...a region'svegetationis its plantlifewhichconsistsof speciesof theflorafound in varying quantity and size, some prominent,others scattered and merging into the ... To reach a thorough understandingof vegetation the flora must first be background understood,but the flora may be studiedwithout an exact and completeknowledgeof the vegetation.Thus a region'sfloraand its vegetationare two quitedifferentthingswhichshould not be confused:the firstmeansparticularly the numbersof the distinctplantspecieswhichone observes,thesecondtheirproportionsandassociations... thevegetationmaybe luxuriantandthe florapoor, the soil sterileand the florarich."8 A keen interest in the phenomena of vegetation is displayed within the advanced and wide-ranging work of the French botanist Henri Lecoq, Professor of Natural History at Clermont-Ferrand.59Lecoq made his allegiance to Humboldt's programme for plant geography quite clear: 'Mais alors un livre me tomba sous la main; j'avais appris, quoique bien jeune encore, a respecter le nom de son auteur, et ce livre, en me revelant une science que je soupconnais sans la connaltre, mit de l'ordre dans mes idees; et dirigea par la suite une partie de mes etudes; c'etait L'Essai de ge'ographiebotanique du celebre Alexandre de

Humboldt.'60 56 0. Heer, Beitrage zur Pflanzengeographie,Zurich, 1835, 1. The last sentence quoted is from a footnote on the same page. 57 For discussions of Linnaeus's 'ecological' writings, see, for example, G. E. Du Rietz, 'Linneaus as a phytogeographer', Vegetatio (1957), 7, 161-8; F. N. Egerton, 'Changing concepts of the balance of nature', QuarterlyReview of Biology (1973), 48,322-50; R. Stauffer,'Ecology in the long manuscriptof Darwin's " Origin of Species" and Linnaeus's "Oeconomy of Nature"', Proceedingsof the American Philosophical Society (1960), 104, 235-41. For an account of Wahlenberg's distinctive programme of researches, see T. Soderqvist, The Ecologists: From Merry Naturalists to Saviours of the Nation, Stockholm, 1986, 20. The Ecologists is a valuable source for the history of Scandinavian natural history and ecology generally. 58 J. Thurmann, Essai de phytostatique applique'ea'la chaine du Jura et aux contre'esvoisines, Berne, 1849, 22, my translation. Thurmann is also briefly discussed by A. G. Duff, 'The Institutionalisation of Ecology in Britain and the United States, 1890-1918', Ph.D. thesis, University of Manchester, 1980, 26, and by Drouin, op. cit. (7),

64-5. 59 For biographical details of Lecoq, see M. Chassagne, 'Le professeur Henri Lecoq, 1807-71', Bulletin de la Societe'Botanique de France (1928), 75, 662-70. 60 H. Lecoq, Etudes sur la gedographiebotanique de l'Europe, 9 vols., Paris, 1854, i, p. v.

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Lecoqworkedwith a distinctionbetweenthe studyof floraand the studyof vegetation whichwas verysimilarto Thurmann's:'IIy a doncunetres-grandediff6renceentrela flore et le tapisvegdtald'unecontree.Lapremierefournitles materiauxqui serventa constituer le second'.61 Floristicswas to be the servant,not the master,of plant geography: j'aid'uautantquepossible,conserverles anciennesespeces,en les considerant,au besoin,comme des groupes.Je n'ai pas l'intentionde publierune flore ni de discuterdes caracteres,mais seulementde m'occuperde la geographieet de la dispersiondes especesdu plateaucentralde la France.Ce n'estdonc ni par ignorancedes Ecritspublies,ni par negligenceou mauvaisvouloir, queje n'adoptepas la majeurepartiedesespecesnouvelles;j'apprecietout le meritedesbotanistes qui se livrenta cetteetude,mais,pourdes travauxde geographiebotanique,je suis forcede me contentersouventdes groupesau lieu d'especesbien d6finies.62

In his enormoustreatiseon the vegetationof centraland southernFrance,publishedin 1854,Lecoqdescribedmanydifferentvegetationtypes.The distributionof the associations was correlatedwith climatic and environmentalfactors such as the intensityof solar radiation, temperature,soil water content, and so on. In an importanttheoretical discussion,Lecoqclarifiedthe distinctionbetween'sociabilite"(manyplantsof the same specieslivingtogether)and 'association'(manyplantsof differentspecieslivingtogether), thus helpingto codify the work of Humboldtand the earlierHumboldtians.63 Anotherfeatureof Lecoq'sworkwhichidentifieshimwith Humboldtis the significance he gives to the aestheticappreciationof vegetation:'Ce qui frappele voyageurquandil penetrepourla premierefois dans une contree,c'est l'aspectgeneraldu pays; ce sont les sites divers avec leurs caracteressi varies,et l'ensemblequi en resulte.Ces premieres impressionsse transformenten souvenirsqui vous retracenta chaqueinstantles grands tableaux de la nature, mais il est bien rare que l'on se rende compte des causes du pittoresqueet des detailsqui entrentdans ces scenesimposanteset majestueuses.'64 It must be admittedthat not every botanistwho explicitlyacknowledgeda debt to Humboldtwas a Humboldtianplantgeographerin the sensein whichthat designationis employedin this paper.Whatdistinguishedthe practiceof Humboldtianplantgeography from that of floristicplant geographywas not merelyadmirationof Humboldtbut a concernwith vegetationas an autonomousobjectof inquiry.For instance,a passagein praise of Humboldt, very similar to that quoted above from Lecoq, occurs in the introductionto Alphonsede Candolle'sgreatbook, Ge'ograpbie botaniqueraisonne'e.65 De Candolle also correlatedplant distributionwith climatic, physical and geographical factors.However,throughoutthe book he is concernedwith the distributionof species, generaandfamilies,hardlyat all with vegetationtypes.It is significant,for instance,that the illustrationfolded into the firstvolumeof Ge'ographie botaniqueraisonne'eis not a 61 Lecoq, op. cit. (60), i, 7. 62 Lecoq, op. cit. (60), i, p. vii. This passage resonates with later debates about the relative merits of floristic criteria in the classification of vegetation; see, for example, M. Nicolson, 'National styles, divergent classifications: a comparative case study from the history of French and American plant ecology', in Knowledge and Society: Studies in Sociology of Science, Past and Present (ed. L. Hargens, R. A. Jones and A. R. Pickering), Greenwich, CT, 1989, 8, 139-86. 63 Lecoq, op. cit. (60), iv, 58-90. 64 Lecoq, op. cit. (60), i, 134; see also Drouin, op. cit. (7), 71-2. 65 A. de Candolle, Ge'ographiebotanique raisonne'e:Ou exposition des faits principaux et des lois concernant la distribution ge'ographiquedes plantes de l'epoque actuelle, 2 vols., Paris, 1855, i, p. v.

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Humboldtian panorama of natural regions and vegetation types, but a map of the northerly limit of various individual plant species in Europe. De Candolle recognized the existence of what he called Topographie botanique, which he defined, partly, as: 'rechercherquels sont les caracteresde la vegetation des marais, des prairies, des f6rets, etc., dans quelles proportions les diverses categories de plantes s'y trouvent representees,par quels motifs certaines espees en sont exclues, etc.; ce serait une Topographie botanique'.66 But he was not very interested in pursuing that study: 'Cette distribution locale, ou topographie des plantes, pourrait constituer une branche de la science, moins importante sans doute que la geographie botanique, mais offrant des developpements analogues ... Mon intention n'est pas d'en parler ici fort en detail, car se serait sortir de mon sujet.'67 De Candolle personified the floristic side of the dichotomy which existed in nineteenthcentury plant geography. But the fact that de Candolle felt obliged to damn topographical botany with faint praise in this way must surely indicate that Humboldtian practitioners were having a substantial impact within the practice of botany and plant geography as a whole by this time. NORTHERN

HUMBOLDTIANS

- VON POST AND HULT

In the early work of the Swedish botanist, Hampus von Post, the emphasis was firmly placed upon vegetational rather than floristic plant geography. To von Post, the major challenge facing plant geographers was the investigation of 'those associations of several plant species which together occupy a similar place... on the earth's surface'.68Von Post urged his fellow Scandinavian botanists to adopt the physiognomic approach and advocated the study of such matters as the interactions between the different species of plants within the association. In his descriptions of the vegetation of Sweden, von Post organized individual plots into associations and then classified these local associations into vegetationsgrupper,a category roughly corresponding in status, if not in size or scope, to the 'formation' of Grisebach and other Humboldtian botanists.69 Of the later ScandinavianHumboldtians, the work of the Finnish botanist Ragnar Hult is particularly interesting. In his important article, 'Forsok till analytisk behandling av vaxtformationerna' (Attemptto make an analytical scheme of plant formations), published in 1881, Hult unequivocally identified himself as working within a tradition of vegetation science that was distinctively independent of floristics or systematics: 66 De Candolle, op. cit. (65), i, 419f; see also ii, 1175-6. 67 De Candolle, op. cit. (65), i, 419. Ronald Tobey has argued that de Candolle denied the possibility of the study of vegetation. But, as this passage indicates, de Candolle acknowledged the existence of a distinctive vegetation science, while himself following another tradition of botanical practice. See R. C. Tobey, Saving the Prairies: The Life Cycle of the Founding School of American Plant Ecology, 1895-1955, Berkeley, 1981, 100. For further comment on Tobey's account of the origins of plant ecology, see M. Nicolson, 'No longer a stranger?A decade in the history of ecology', History of Science (1988), 26, 183-200. 68 H. von Post, Forsok till en systematisk uppstallning av vexstallena i mellersta Sverige, Stockholm, 1862. I am greatly indebted to Sigridur Oladottir for making this text accessible to me and for providing me with a translation of the passages quoted from Ragnar Hult. For an account of von Post, see Soderqvist, op. cit. (57), 24-9. 69 See Whittaker, op. cit. (25), 23.

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When botanical geography first appeared, it was an appendage and auxiliary to systematic botany. Species, genus, families and higher taxa were studied from the point of view of site conditions, distribution and mode of distribution and from this researchconclusions were drawn regarding each taxon's habitat and so on. It was ignored or overlooked that there also existed other individually very obvious groups of plants as well as the systematic taxa, 'one did not see the wood for the trees'."7

Hult creditedHumboldtwith having been the first to recognizethe existenceof the collectivegroupingsof plants,and with havingcreateda vegetationalplantgeography: It needed Humboldt's mighty spirit... to discover these groups and to realize their immense significance in the geography and history of the Earth. In his travels through tropical America he was struck by the definite outlook which certain characteristic and dominant plants lent the landscape. He saw the shifting expression of this outlook in the dark of the jungle and on the monotonous plains of the savannas, on the palm decked tidal beaches, and in the treeless high mountains of the Andes; he saw the vegetation of all the regions together within a limited space, but organized according to a system of law which he could not fail to notice ... he emphasised primarily the physiognomy and the growth forms of plants, the interdependenceof these features from systematic relations, and their role in the characterof the landscape, as well as these groups' mode of growing together, the monotony of the colonies of the social plants in contrast to the diversity of forms and variations in the tropical jungle.7'

EvidentlyHult also sharedHumboldt'sconcernthatthe plantgeographershouldcultivate an aesthetic,as well as a scientific,responseto vegetation.In this article,Hult tracedthe history of the study of the types of vegetation,from its origins with Alexandervon Humboldt,throughthe work of Grisebachand Kernervon Marilaun,to his own time. He developeda systemin whichbothlifeHult'sapproachwas strictlyphysiognomic.72 formsandthe vegetationunitsbaseduponthemwerenarrowlydefined,distancinghimself from those plant geographerswho had followed more closely the exampleof Schouw, whose approachto plantcommunitieshe regardedas 'deductive': Because if one is to go for example to a moor in the middle of Finland, one can see there in an area where no differencesin the chemical or physical conditions can be shown at least two sharply divided plant groupings alternating in patches. One is an even and dense mass of Clandina silvatica, with other lichens sprinkled in, as well as Polytricba and low Empetrum; the other is a similarly thick and even mat of Calluna vulgaris, with a sparse undergrowth of Cladonia, Hylcomia and Polytricha, as well as ... a few other plants. Here we can thus see an intimate mixture on the same location of two plant communities, which are in sharp contrast to each other. And these have to be united by the deductive school of thought, which according to the varying degrees of dampness of the soil, distinguishes formations with barely distinguishablevegetation.73

The defectsof the 'deductiveschoolof thought'flowed,in Hult'sopinion,fromits being principallyconcernedwith the physical environmentrather than primarilywith the vegetationitself. Hult's developmentof the Humboldtianlegacy was, thus, selective. He adopted physiognomicanalysisand the characterizationof vegetationunits, for example, but neglectedenvironmetry. By the secondhalf of the nineteenthcentury,Humboldtianplant 70 R. Hult, 'F6rs6k till analytisk behandling av vaxtformationerna', Societas pro Fauna et Flora Fennica Meddelanden (1881), 8, 1-155, on 1. See note 68 above. 71 Hult, op. cit. (70), on 1. 72 Whittaker, op. cit. (25), 23-4. 73 Hult, op. cit. (70), 9.

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geography was undergoingprocesses of internaldivergencein the course of its development as a research tradition. Following the nature of these divergences is essential if the role played by Humboldtian plant geography within the development of plant ecology is to be accurately assessed. For example, in Hult's accounts of Swedish vegetation, we can discern emphases which were to become quite characteristic of the Uppsala school of phytosociology in the twentieth century - the recognition of vegetation units by autonomous vegetational criteria rather than by environmental factors, and a preoccupation with quite small units of plant cover.74

THE LINK TO A 'SELF-CONSCIOUS'

ECOLOGY

It has often been argued that plant ecology sprung quite suddenly into existence at the end of the nineteenth century - fathered by the inspiration of Warming's, Drude's and Schimper's great texts, respectively Lehrbuch der okologischen Pflanzen-geographie, published in 1896 (originally in Danish in 1895), Deutscblands Pflanzengeograpbie,1896, and Pflanzengeographie auf physiologischer Grundlage, 1898.75 The reminiscences of British and American ecologists often tell how one or other of these books so excited the readerthat he set out to put the precepts of the Continental author into practice in his own country.76No doubt these accounts are true, to an extent.77Such stories have, however, become received as an explanation of the origin of the subject, rather than its transmission to Britain or America. As Worster put it:'Their [Warming,Drude, Schimper]work during the 1890s transformedOecologie from just another neologism to a functioning science with its own peculiar hold on reality'." Goodland dubbed Warming 'the founder of plant ecology .79 However, Harry Godwin has pointed out the obvious oversimplificationinherent in this account of the origin of plant ecology.80 He traced the influential Zurich-Montpellier school of phytosociology back beyond 1890 and showed that, by this time, one of its founders, Carl Schroter, was well established on a course of ecological researches, quite independent of Warming or Schimper. Likewise, the Swedish botanist Rutger Sernander, 74 See C. E. Moss, 'The fundamental unit of vegetation', New Phytologist (1910), 9, 18-53; and Whittaker, op. cit. (25). 75 E. Warming, Lehrbuch der okologischen Pflanzen-geographie: Eine Einfuhrung in die Kenntnis der Pflanzenvereine,Berlin, 1896; 0. Drude, Deutschlands Pflanzengeographie:Ein geographischesCharakterbildder Flora von Deutschland und den angrenzenden Alpen-Sowie Karparthenldndern,Stuttgart, 1896; A. F. W. Schimper, Pflanzengeographie auf physiologischer Grundlage, Jena, 1898. Warming's text was originally published in Danish as Plantesamfund: Grundtraekaf den 0kologiske plantegeografi, Copenhagen, 1895. 76 A. G. Tansley, 'The early history of modern plant ecology', Journal of Ecology (1947), 35, 130-7; D. Worster, Nature's Economy: A History of Ecological Ideas, Cambridge, 1985, 207; C. Adams and G. D. Fuller, 'Henry Chandler Cowles, physiographic plant ecologist', Annals of the Association of American Geographers (1940), 30, 39-43. 77 For a discussion of the nature of H. C. Cowles's debt to Warming, see M. Nicolson, 'Henry Allan Gleason and the individualistichypothesis of the plant community: the structureof a botanist's career', Botanical Review (1990), 56, 91-161, on 97-9. 78 Worster, op. cit. (76), 198. 79 R. J. Goodland, 'The tropical origin of ecology: EugenWarming'sjubilee', Oikos (1975), 26,240-5, on 241. 80 H. Godwin, 'Sir Arthur Tansley: the man and the subject', Journal of Ecology (1977), 65, 1-26 on 8.

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underwhose leadershipthe Uppsalaschool was founded,had alreadystartedapplying quantitativemethodsto the studyof vegetationby the mid-1880s.Goodwinthus argued that the two principalContinentalschools of vegetationscience were foundedbefore Warmingand Schimperpublishedtheir majortexts. The existenceof plant ecology, or somethingvery like it, mustbe the explanationfor Warmingand Schimper,and not vice versa.Indeed,it only requiresthe briefestlook at the LehrbuchderdkologischenPflanzengeographieandthe Pflanzengeograpbie aufphysiologiscber Grundlage- at theirsize,scope - to see that theycouldnot havebeenproducedby men and bibliographiccomprehension who were merelythe rude forefathersof a nascentscientificspecialty.It is evidentthat thesetexts representa culminationas well as a startingpoint. They containthe fruitsof manyyearsof inquiry,of a long traditionof researchinto vegetation. This paperhas describedthe developmentof that tradition,extendingfromHumboldt throughGrisebach,Heer,Hultandthe otherHumboldtians.Onwardconnectionsbetween Humboldtianplantgeographyandtwentieth-century vegetationsciencecan also be readily traced. Carl Schr6ter,for example, was a doctoral student of Oswald Heer at the Universityof Zurich.He was a sufficiently favouredstudentto inheritHeer'sprofessorship uponhis deathin 1883.Schroter'sstudentand biographer,EduardRubel,has stressedthe influenceof Kernervon Marilaun'sPflanzenleben der DonaulanderuponSchroter'searly phytogeographicalwork.81 As its name suggests, Kerner'sPflanzenlebenprovidedan importantmodel for Schroter'sown Das Pflanzenlebender Alpe. Thus we see that the Zurich-Montpellierschool, which was active throughoutthe twentiethcentury,had, throughits founders,directlinksto the earlyHumboldtians.82 The samemay be said of its northerncounterpartin Uppsala.RutgerSernanderwas an assistantto RagnarHultin the early1880s.The mostfamousnamesof the Uppsalaschool - Du Rietz, Osvald, Nordhagenand L. von Post- were all studentsand disciplesof A direct connectionis again evident between the practiceand usage of Sernander.83 Humboldtianplant geographyand the 'self-conscious'ecologyor plant sociologyof the twentiethcentury. A similarstory may be told of OscarDrude,one of the trio namedby Worsteras the fathersof ecology.Drudewas, from1871to 1873,researchassistantto AugustGrisebach at Gottingen.84He explicitly presentedhis later researchas continuingGrisebach's LikeGrisebach,Drudeput a strongemphasison the investigationsin plantgeography.85 unitaryintegrityof the regionalformation,the characterof whichwas determinedby the regionalclimate.Formationswere,however,internallyheterogeneousowing to the effect of topographyupon the vegetation.Drude furtherdevelopedGrisebach'ssystem of growth-forms.However, he characterizedthe formationrathermore floristicallythan Grisebachhad done, recognizingsmallersubsidiaryunits,identifiableby locallydominant 81 E. Rubel, 'Uber die Entwicklung der Gesellschaftmorphologie', Journal of Ecology (1921), 8, 18-40; see also idem, 'Ecology, plant geography and geobotany: their history and aim', Botanical Gazette (1927), 84,428-39. 82 See also R. W. Becking, 'The Zurich-Montpellier school of phytosociology', Botanical Review (1957), 23, 412-88. 83 Godwin, op. cit. (80), 8. See also Soderqvist, op. cit. (57), 87-9. 84 R. Zaunick, 'Drude, Carl Georg Oscar, Botaniker', Neue Deutsche Biographie, iv, 138. 85 0. Drude's Handbuch der Pflanzengeographie,Stuttgart, 1890, is dedicated to Grisebach; see also p. ix.

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species, which he termed 'Bestdnde'. Drude's Deutschlands Pflanzengeographie was adopted by Roscoe Pound and FredericClements as the model for their early investigations into the plant geography of Nebraska.86Clements went on to found one of the most important schools of plant ecology in America.87 Several of the principal themes of Schimper's Plant Geography upon a Physiological Basis, to give it its English title, have been identified above as distinctive concerns of the Humboldtian tradition. Schimper structured his treatment of the world's plant cover around a classification of vegetation into regions, formations and smaller units. This system was, at least largely, physiognomic: Climaticformationsmay be tracedbackto threechieftypes- woodland,grassland,and desert. Woodlandis constitutedessentiallyof woodyplants,andis termedforestif treesgrowin a closed condition;bushwoodwhen shrubsare so abundantas to keep the crownsof the trees from touchingone another;shrubwoodwhereshrubsconstitutethe chieffeature.88 Schimper also sought to correlate the occurrence of vegetation types with the physical environment. His book is replete with tables of mean temperaturesin the Brazilianforests, humidity in the Antarctic, the variation of day-lengthwith latitude - all mannerof physical data from all the major vegetational regions of the world. Schimper further developed Grisebach's distinction between different means of environmental control: - theclimaticor districtformations, two oecologicalgroupsof formationsshouldbedistinguished the characterof whosevegetationis governedby atmosphericprecipitations, andthe edaphicor local formationswhose vegetationis chieflydeterminedby the natureof the soil.89 Folded in at the end of the book is a set of maps showing the distribution of the world's rainfall, correlated with the occurrence of the climatic vegetation-types. There are, of course, many novel elements in Schimper's work. German botany had undergone many changes in the later decades of the nineteenth century. With the development of plant morphology and physiology, plant science had become more experimentally oriented and less holistic. The problem of adaptation, often conceived anew in Darwinian terms, altered the character of studies into the external and internal structureof plants.90The aesthetic appreciation of plants and vegetation had become less of an explicit concern, at least within the scientific literature. However, the new trends in botanical science did not completely swamp the older programme of Humboldtian plant geography. Rather they interacted with it to produce the new style of investigation which was to be called 'plant ecology'. The publication of Grisebach's Die Vegetation der Erde stimulated many botanists to study the relationship between vegetation and environment. For instance, the anatomist 86 R. Pound and F. E. Clements, The Phytogeographyof Nebraska, Lincoln, NE, 1900. 87 J. B. Hagen, An Entangled Bank: The Origins of Ecosystem Ecology, New Brunswick, 1992; Nicolson, op. cit. (62); Tobey, op. cit. (67). 88 A. F. W. Schimper, Plant Geography upon a Physiological Basis (tr. W. R. Fisher, ed. P. Groom and I. Bayley Balfour), Oxford, 1903, 162. 89 Schimper, op. cit. (88), 161. For a detailed comparison of the work of Schimper and Grisebach, see E. Cittadino, Nature as the Laboratory:Darwinian Plant Ecology in the GermanEmpire, 1880-1900, Cambridge, 1990, 112-15. 90 This point is well made by Cittadino, op. cit. (89).

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Schwendener assignedto a doctoralstudentthe taskof determiningwhetherthe vegetation zones describedby Grisebachwere reflectedin anatomicaldifferencesbetween plant But,generally,those who appliedthe species.91Grisebachwas staunchlyanti-Darwinian. fruitsof the new studiesof physiologyand morphologyto understandingthe life of the plantin the wild interpretedstructureandgrowth-formin Darwinianterms.Thus,in the decades after Grisebach,vegetationalplant geographyincorporatedphysiology,morphologyand Darwinisminto its explanatorystructure.Schimper'swork well exemplifies thesenew developments.Schimperwas, however,thoroughlyacquaintedwith Grisebach's work and madefrequentreferencesto his majorpublication: The connexionbetweenthe formsof plantsandthe externalconditionsat differentpointson the of oecologicalplant-geography, whichhas only recently earth'ssurfaceformsthe subject-matter becomea prominentsubjectof interest,althoughit founda placein earlierworks,especiallyin Grisebach'svaluableVegetation der Erde.92 Schimper's classification of the world's vegetation was very similar to Grisebach's. But Schimper was a committed Darwinian and interpretedthe facts of plant distribution and morphology accordingly. As Cittadino has put it: 'From the viewpoint of plant geography, Schimper'swork falls within the tradition of Grisebachbut ... Schimperwas concerned not only with the nature but also with the origin of adaptation'.93Likewise it was Schimper's programmatic intention to incorporate new physiological interpretations into his analysis of the correlation between vegetation and environment: 'The oecology of plantdistribution will succeed in opening out new paths on condition only that it leans closely on experimentalphysiology, for it presupposes an accurate knowledge of the condition of the life of plants which experiment alone can bestow'. It is, thus, impossible to claim that Schimper's 'oecological plant-geography' was a simple continuation or articulation of Humboldtian plant geography. But the content of Plant Geography upon a Physiological Basis was nevertheless strongly conditioned by the prior existence of the Humboldtian tradition of inquiry. We can see a similar relationship between new and old elements in the work of Eugen Warming, Professor of Botany at the University of Copenhagen, particularly in Oecology of Plants: An Introduction to the Study of Plant Communities. In terms reminiscent of Franz Meyen, Warming distinguished his research activity from floristic plant geography which 'is concerned with... the compilation of a 'Flora', that is, a list of species growing within a larger or smaller area': has entirelydifferentendsin view: Oecologicalplant-geography 1. To findout whichspeciesare commonlyassociatedtogether... 2. To sketchthe physiognomyof the vegetationand the landscape... 3. To answerthe questionsWhyeach specieshas its own specialhabitand habitat? Why the species congregate to form definite communities? Why these have a characteristicphysiognomy? 91 Cittadino, op. cit. (89), 60. 92 Schimper, op. cit. (88), p. vi. 93 E. Cittadino, 'Plant Adaption and Natural Selection after Darwin: Physiological Plant Ecology 1880-1900', Ph.D. thesis, University of Wisconsin, Madison, 1981, 151. 94 Schimper, op. cit. (88), p. vi.

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4. To investigatethe problemsconcerningthe economy of plants... We thus come to the of plants.96 considerationof the Growth-forms Again, there is much here that is familiar - the concern with physiognomy and 'definite communities', in particular. Apart from morphology, the major novel element in Warming's text was an interest in evolutionary adaptation. But this new concern was expressed within the established framework of vegetational research. Warming continued to classify vegetation into natural units, according to physiognomy. He elaborated a new classification system for growth-forms - contributing to a form of research practice he explicitly traced from Humboldt through Grisebach to his own contemporary Oscar Drude. Warmingwas concernedwith relatively smaller, more localized pieces of vegetation than Schimper'sclimatic formations. But he was similarlyconcernedto correlatethese with the physical environment and to understandthe effect of the environment on the structure of the plant. Warming was not, however, a strict natural selectionist. He held that both species and physiognomic growth-forms 'stand in perfect harmony with the environment' and that 'plants possess a peculiar inherent... faculty by the exercise of which they directly adapt themselves' to their surroundings.96In other words, vegetation is the creation and the expression of the environment- a very Humboldtian conception.97 Warming's plant geography, like Schimper's, possessed much of the character it did because Alexander von Humboldt and the Humboldtians had worked before him. It is noteworthy that the bibliography of Oecology of Plants reads like a roll-call of Humboldtian plant geography- Hult, Schouw, Kerner von Marilaun, Meyen - back to Humboldt himself. And, most significantly, neither Warming nor Schimper cited any author who wrote prior to Humboldt. No eighteenth-centurybotanist, not even Linnaeus, no one who worked within the old episteme, is cited.98The Essai sur la geographie de plantes is a principal startingpoint for Warming's and Schimper'sresearchenterprise- an enterprise which had become, by the time of the publication of their major texts, unequivocally 'plant ecology'. 95 E. Warming, Oecology of Plants: An Introduction to the Study of Plant Communities, Oxford, 1909, 2-3, emphasis in the original. 96 Tobey, op. cit. (67), 104 claims that the English translation 'introduced a Humboldtian nuance into [Warming's] Darwinian theory'. Thus, Tobey argues, it cannot be said that Warming supported the idea of definite communities in the Humboldtian sense. But see 0. Muller, 'Warming, Eugen', DSB, xiv, 181-2, for evidence to the contrary, drawn from the original Danish Plantesamfund,Warming, op. cit. (75). For instances of Warming employing the concept of the plant community in the characterizationof vegetation in the field, see E. Warming, 'Om Gr0nlands vegetation', Meddelanden om Gr0nland (1888), 12, 1-245 (Frenchsummary). See also note 67 above. 97 This would probably also have been Grisebach's view on the origin of plant form. Tobey has argued that Warming's 'Darwinian' concept of a competitive struggle for survival distinguishes him from the 'idealistic' tradition of Humboldt and Drude. Warming was certainly interested in evolution and the origin of adaptation. But he, unlike Schimper, was not a Darwinian in the sense that he favoured the mechanism of natural selection. In fact, like many early ecologists, he was famous for his eclectic Neo-Lamarckianism.The Humboldtian research tradition was sufficientlydiverse to allow plant communities to be conceived of in a variety of differentways but there is no evidence in Warming's work that he did not allow the existence of, to use Tobey's terminology, 'functioning communities of ontological status'. See also notes 67 and 96 above. 98 For the identification of Humboldt with epistemic change in Michel Foucault's sense, see Nicolson, op. cit. (3); also M. Nicolson, 'Was there a Linneanecology?', unpublishedpaper, copies available from the author; and Acot, op. cit. (7).

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CONCLUSIONS As Susan Cannon has made very clear, Alexander von Humboldt's complex of interests does not fit neatly within twentieth-century disciplinary boundaries.99 It would be downright misleading to refer to Humboldt as an ecologist; one even calls him a plant geographer at some risk of faulty and anachronistic identification. Humboldt was, one might say, primarily a 'Kosmos-ologist';100 or, to use his own term, a student of 'la physique generale' - a grand programme of inquiry within which plant geography had a key role. Plant geography was, to Humboldt, a most important synthetic subject - a study indeed of 'Kosmos-ological ' significance. It was thus quite different in scope and importance from what we now know as plant geography or, even, ecology. However, it remains legitimate to trace within Humboldt's work the roots of concepts and practical exemplars later to be employed within the cognitive frameworks of more specific disciplines. Not all the Humboldtians could be as Kosmos-ological or as polymathic as the great man himself. The scientific enterprise was, moreover, even in Humboldt's lifetime, increasingly becoming more rigidly demarcated into narrower specialties.?1' Later researchers- Humboldtians in that they were more or less directly associated with Humboldt, shared many of his concerns, articulated the exemplars contained in his scientific writings, used instruments in the manner he recommended, studied some of the objects he studied, and so forth - employed the resources presented to them by his work in more specialized, more circumscribedcontexts, which more closely resemble our present disciplinary divisions. Grisebach and Hult, for example, were essentially professional botanists - they were not necessarily students of all the ramifications of 'la physique generale'. But scientists might be more restrictedin their range of interests than Humboldt was and yet still be practitionersof Humboldtian science, as they saw it applicable within their own subject areas. As a research programme Humboldtian plant geography was certainly flexible; it diverged into several schools; it incorporated new elements. But, with its distinctive emphasis upon the study of the collective phenomena of vegetation, it is discernible throughout the nineteenth century nevertheless. The vegetational tradition of plant geography extended into an age of greater specialization in natural science - extended indeed to the birth of ecology as an identifiable specialty in its own right. As acknowledged above, the twentieth-century scientific specialty of plant ecology had more than one precursor.McIntosh has written of 'self-conscious' ecology emerging from a 'complex interaction between 19th-century natural history and 19th-century physiology'. 102 In the light of the work of Cittadino and Coleman, evolutionary studies should be added to the list of historical antecedents.'03Drouin has intriguingly suggested that it 99 Cannon, op. cit. (11). 100 The allusion is to the title of Humboldt's most ambitious text: A. von Humboldt, Kosmos: Entwurf einer physischen Weltbeschreibung,S vols., Stuttgart and Tuebingen, 1845-62. 101 Some of the consequences of this changing context for Humboldtian science have been explored by Cannon, op. cit. (11). 102 R. P. McIntosh, The Background of Ecology: Concept and Theory, Cambridge, 1985, 26. 103 Cittadino, op. cit. (89); W. Coleman, 'Evolution into ecology? The strategy of Warming's oecological plant geography', Journal of the History of Biology (1986), 19, 181-96.

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might be worthwhile investigating whether Continental floristic plant geography, as developed by Alphonse de Candolle, also played an importantpart.'04This paper contends that the recognition of the existence, and distinctiveness,of a Humboldtian programmefor the geography of vegetation is, likewise, a valuable aid to the analysis of the undoubtedly complicated and heterogenous origins of the discipline. A number of first-generation ecologists, such as Schroter, Sernander, Drude, Warming and Schimper, must also be counted as third or fourth generation students of Humboldtian plant geography. Thus, the vegetational geography of Schouw, Grisebach, Kerner von Marilaun, Lecoq and their peers, was one of the roots from which modern ecology sprung.

104 Drouin, op. cit. (7), 73-80.

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