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History of Plant Ecology

Malcolm Nicolson, University of Glasgow, Glasgow, UK
Plant ecology is the study of the collective phenomena of plants. It dates from the nineteenth century, having its origins in Humboldtian plant geography and in Darwinian physiological anatomy. Much important work was done in the USA and Britain, both before and shortly after World War II. The study of plant communities is now a major aspect of ecosystem ecology.

Introductory article
Article Contents
. Introduction . Humboldtian Plant Geography . Physiological Anatomy . Plant Ecology in America and Britain . New Ecology

Introduction
The use of the term ‘plant ecology’, as the accepted name of a distinctive form of scientific inquiry, dates from the 1890s. But the history of the discipline extends considerably further back into the nineteenth century. Plant ecology has traditionally had two facets – the study of how, and why, different plant species grow together in communities and the study of the relations of individual plant species to their environment. The former branch, known as ‘synecology’ or ‘vegetation science’, is both the more dominant and the older partner, having its historical roots in the plant geography of Alexander von Humboldt. The latter branch, known as ‘autoecology’ or ‘physiological plant ecology’, derives substantially from the impact of Darwinism upon mid-nineteenth-century plant anatomy and physiology.

example, an association in which one of the species of the oak, Quercus, was the most numerous tree, was termed a Quercetum. August Grisebach further developed Humboldt’s system of classification by growth forms and coined the term ‘formation’ for a type of vegetation characterized solely by physiognomic criteria. In 1872, Grisebach published his great book, Die vegetation der Erde (The Vegetation of the Earth), a comprehensive treatise of physiognomic plant geography. By the 1880s, other plant geographers, in France, Switzerland and Scandinavia, had developed alternative systems of vegetational classification based upon more floristic criteria.

Physiological Anatomy
Following the appearance of Darwin’s On the Origin of Species in 1859, a number of plant physiologists, notably Simon Schwender at the University of Berlin, rebelled against the strict division between morphology and physiology which characterized German botany at that time and developed a programme of research in which a systematic attempt was made to correlate morphological structure with adaptive function, in the context of the life of the plant in the wild. Upon reading Die Vegetation der Erde, Schwender realized that his ‘physiological anatomy’, as it was termed, could potentially be applied to the investigation of the geographical distribution of plants and vegetation types. The challenge was to elucidate, from a physiological point of view, how particular plants were adapted to particular habitats. A similar research programme was developed simultaneously by Anton Schimper, a morphologist by training. German botanists were also inspired to elucidate the relationship between plants and the environment by their experience of exotic tropical vegetation. In the last decade of the nineteenth century, two major books were published– Pflanzengeographie auf physiologischer Grundlage by Schimper and Lehrbuch der o¨kologishen Pflanzen-geographie by Eugen Warming. Both books drew jointly upon Humboldtian plant geography and physiological anatomy to provide a comprehensive
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Humboldtian Plant Geography
Up until the end of the eighteenth century, botanists concentrated their efforts on finding individual plants and classifying the specimens into species and higher taxa. In 1790, however, Alexander von Humboldt set out a programme for a new form of plant geography which ‘traces the connections and relations by which all plants are bound together among themselves, designates in what lands they are found, in what atmospheric conditions they live ...’. Humboldt drew attention to the fact that plants tended to grow together in recognizable and recurring groups and was the first to use the term ‘association’, albeit informally, to describe these types of vegetation. He developed a non-floristic classification of vegetation types, based upon the growth form of the constituent plants. Using a number of measuring instruments, Humboldt sought to correlate the geographical distribution of vegetation with a wide variety of environmental variables. Humboldt’s field work was done mainly in South America but later investigators, Anton Kerner for example, employed and refined his methods in the investigation of European vegetation. In 1822, Joachim Schouw devised the ‘-etum’ nomenclature, whereby, for

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trained in floristics but aware of the decreasing fashionability of pure natural history. European examplars were also quickly adopted in the University of Nebraska. it promised a privileged insight into the history of vegetation for:In many cases . Cowles called his new approach ‘physiographic ecology’. It seemed to uncover the laws of vegetational development. his student and assistant. Cowles. Minneapolis. and then Henry Cowles. academic plant ecology developed significant links with agronomy. Tansley founded the British Vegetation Committee. in 1913. His methods were rapidly applied throughout North America by his students. 1901]. he was interested in the application of ecological knowledge to the practical problems of farming and ranching on the Great Plains. F. The University of Nebraska was a LandGrant college and. in keeping with its institutional ethos. the latter as Oecology of Plants: An introduction to the study of plant communities (1909). how types of vegetation replaced one another over time. and into the relation of its constituent species to environmental factors. adopted Drude’s work as the model for their pioneering study. In 1899. cleverly meshed the European systems of vegetational classification with the theory of geomorphological base-levelling which had been developed by American geologists such as his teacher. under whose auspices the valuable ‘guide book’ Types of British Vegetation was published in 1911. Warming had based his classifications upon specific habitat factors such as the water content of the soil. in Minnesota. Arthur Tansley. species composition and distribution. Thus by. the Germaninspired shift in emphasis from floristics and taxomony as the dominant modes of botanical instruction and inquiry towards the study of plants by means of microscopy and experimentation. namely topography. where Charles Bessey was Professor of Botany. which he and Pound had devised. Professor of Botany at University College London. moreover. there is a horizontal order of succession at the present time that resembles the vertical [i. which described the quantitative analysis of vegetation by the quadrat and transect techniques. Bessey’s students. He served as a consultant to the Soil Conservation Service and the Great Plains Drouth Committee. in Chicago. laid out in bands parallel with the water’s edge. [HC Cowles. deciduous woodland over much of temperate North America. The Phytogeography of Nebraska. For example. wet and moist soils – the most developed form of vegetation being found in the intermediate conditions.els. In America first Conway MacMillan. In particular. and was one of the most influential teachers of the subject in the United States. Soil water content was chiefly determined by the slope and elevation of the terrain. a student of Grisebach. Roscoe Pound and Frederic Clements. Bessey encouraged his students to pursue research topics that were both scientifically worthwhile and economically useful. Rollin Salisbury. In Clements’ hands. Also in the same decade. Schimper and Warming’s books were quickly translated into English. conveyed a powerful sense both of the ubiquity of vegetational change and of an underlying predictable regularity within that change. Tansley published the definitive The British Islands and their Vegetation in ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www. The phenomena of ‘succession’. translated Kerner’s work and developed the field study of coastal plant communities. building upon this earlier work. Cowles’s theory. Plant Ecology in America and Britain The influence of these texts in English-speaking countries was immense. This form of vegetation. hydrophytic and mesophytic plant communities. horticulture and forestry. Field-orientated botanists. In 1905 Clements produced the textbook Research Methods in Ecology. walking back from a sandy shore one could see. Bessey was the leading American exponent of the so-called ‘New Botany’. furthest from the water. found ecological plant geography to be a fruitful expression of their skills and interests. notably by W. were inspired by Warming’s text to set out to classify their local vegetation along similar lines. W. which culminated in 1916 with the publication of Plant Succession – An Analysis of the Development of Vegetation. as the continental landmass of America was gradually eroding toward the condition of a peneplain. trending toward the most mesophytic vegetation type possible under the regional climate. was dubbed the ‘climatic climax’.e. In Britain. for example. land management. Deutchsland Pflanzengeographie. the temporal] succession of 2 which we now have only the topmost member. the British Ecological Society – the American equivalent dates from two years later. perhaps most excitingly of all. Clements’ research programme was developed further in an impressive series of books and articles. he recognized xerophytic. had long been studied by botanists. And. Cooper became Professor of Ecology at the University of Minnesota. began to teach a course based on Warming’s Lehrbuch.. Oliver. The Vegetation Committee formed the nucleus around which was founded. its character. Cooper at Glacier Bay in Alaska. But Cowles argued that soil conditions were themselves dependent upon a more fundamental environmental variable. Oscar Drude. produced his classic study..History of Plant Ecology outline of what was now called ‘plant ecology’ – a programme of research into vegetation. Thus.net . associated respectively with dry. In 1904. and by the use of recording instruments in the field. S. the development stages of the deciduous forest which was now established upon the stabilized and humuscovered mature sand dunes. the former as Plant Geography upon a Physiological Basis (1903). so its vegetation was becoming more mesophytic in character.

els. Tansley. scarcely even a vegetational unit’ but was merely the product of the ‘co-incidence’ of plants able to live in the same location. Botanical Gazette 31: 73. McIntosh RP (1985) The Background of Ecology: Concept and Theory. a study of the origin. fungi. Cambridge: Cambridge University Press. Henry Allan Gleason. Hagen JB (1992) An Entangled Bank: The Origins of Ecosystem Ecology. the character of biological theorizing was transformed by advances in population genetics and by the development of the Neo-Darwinian synthesis. especially in southern Europe and Scandinavia. bacteria and associated inanimate material should be studied together as components of an interrelated functioning entity.History of Plant Ecology 1939 and was the founding chairman of the Nature Conservancy Council. By the 1960s. Cowles HC (1901) The physiographic ecology of Chicago and vicinity. as developed by plant ecologists. however. The Hague: Junk. while a valid object of study. Darwinian emphasis. as the work of Herbert Bormann and Gene Likens at Hubbard Brook Experimental Forest. Research techniques and equipment were increasingly borrowed from physics and chemistry. larger teams of workers were employed. ordered. Nevertheless. In America. by Robert Whittaker. Much European vegetation science. remained central to much ecosystem theory. coined the term ‘ecosystem’. development and classification of plant societies. however. was ‘not an organism. These natural units of vegetation. indeed ‘super-organismic’. vindicated Gleason’s individualistic hypothesis. The new term conveyed the suggestion that plants. forestry and related fields. the concepts of succession and climax. integration. Detailed quantitative studies of the vegetation of Wisconsin. Cambridge: Cambridge University Press. These researches formed the departure point for the mathematically complex ordination and classification studies which were the cutting edge of vegetation science in the 1960s and early 1970s. Novel modes of investigation. the leading proponent of the community unit theory was Frederic Clements. distinguished the ‘New Ecology’. who claimed to discern in the formation an entity of organic. Similar conclusions were arrived at by British botanists working in the tropical rainforest. became markedly old-fashioned. Throughout the late nineteenth and early twentieth centuries. losing its distinctive disciplinary identity as a more inclusive science of general ecology emerged. British Journal for the History of Science 29: 289–310. and the use of computers for the manipulation of data and the modelling of processes became routine. nearly all vegetation research was predicated upon the conception that plants grow together in definite. In 1935.net 3 . argued that the plant community. New Hampshire exemplifies. albeit now re-expressed with a more adaptive. Clementsian ecology. New Brunswick: Rutgers University Press. were held to constitute the proper subject matter for ecological study – in the same way as plant species were the proper object of study for taxonomic botanists. Worster D (1985) Nature’s Economy: A History of Ecological Ideas. and of the Great Smoky Mountains. repeating communities. such as the study of the flow of nutrients and energy. Further Reading Cittadino E (1990) Nature as the Laboratory: Darwinian Plant Ecology in the German Empire 1800 – 1900. plant ecology was. Whittaker RH (1975) Ordination and Classification of Communities. ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Nature Publishing Group / www. New Ecology After World War II. by John Curtis and his colleagues. By the 1960s. speculative and with a pronounced Lamarckian character. Meanwhile applied plant ecology continues to play a formative role in land management. Nicolson M (1996) Humboldtian plant geography after Humboldt: The link to ecology. ecologists had substantially taken up the challenge of investigating natural environments along these lines. was based around the identification and classification of such units. to a significant extent. animals. wildlife conservation. characterized either by essentially constant physiognomy or essentially constant floristic composition. partly as a corrective to what he regarded as the excesses of Clements’ ‘super-organism’ theory. Cambridge: Cambridge University Press.